ISF 593 

T5 H8 

iCopy 1 



U. S. DEPARTMENT OF AGRICULTURI^ 
BUREAU OF ENTOMOLOGY— BULLETIN No. 72. 

L. 0. HOWARD, Entomologist and Chief of Bureau. 



INFORMATION CONCERNING 



THE NORTH AMERICAN FEVER TICK, 



WITH NOTES ON OTHER SPECIES. 



r.Y 



w. I). hitntp:r and av. a. hooker. 



Issued Novembeb 2, 1907. 




WASHINGTON: 

GOVERNMENT PRINTING OFFI(!E. 

1907. 




aass_^S„FLs9^ 

Book 3jl±1_2 



'i/-. 



U. S. DEPARTMENT OF AGRICULTURE, 

BUREAU OF ENTOMOLOGY- BULLETIN No. 72. 

L. O. HOWARD, Entomologist and Chief of Bureau. 



fl 



INFORMATION CONCERNING 



THE NORTH AMERICAN FEVER TKUv, 



WITH NOTES ON OTHER SPECIES. 



BY 



W. D. HUNTER ani> W. A. HOOKER. 



Issued Novembee 2, 1907. 




WASHINGTON: 

GOVERNMENT PRINTING OFFICE. 

1 \) 7 . 



A 



BUREAU OF EA^TOMOLOar. 

L. (). HowARij, Eniomologist and Chief of Bureau . 

C. L. Marlatt, Eniomologist and Acting Chief in absence of Chief. 

R. S. Cliftox, Chief Clerk. 

F. H. Chittenden', in charge of breeding cxperivicnts. 

A. D. HoPKixs, in charge of forest insect investigations. 

W. D. Hunter, in charge of cotton boll xveevil invest igat ion ,s. 

F. M. Webster, in charge of cereal and forage plant insect irnestigatio7}s. 

A. L. Quaintance, in charge of deciduous fruit insect invent igntions. 

E. F. Phillips, in charge of aviculture. 

D. M. Rogers, in charge of gipsy moth and brown-tail moth imrk. 

A. W. Morrill, engaged in while fly investigations. 

W. F. Fiske, in charge of gipsy moth laboratory. 

W. A. Hooker, engaged in cattle tick life history itirestigationti. 

A. C. Morgan, engaged in tobacco insect investigations. 

R. S. WoGLUM, engaged in hydrocyanic acid gas investigations. 

C. J. GiLLiss, engaged in silk investigations. 

R. P. Currie, assistant in charge of editorial work. 

Mabel Colcord, librarian. 



MAR 31 19C.3 
,D. ot D. 






LHTTHR OF TRANSMITTAL. 



I'. S. Depahtafext of Agkicultihk, 

Bureau of Eintomolocjv, 
WasMngion, I). C, June 19, 1.907. 
Sir: I have the honor to transmit herewith a manuscript prei)ared 
by jNIessrs. W. D. Hunter and W. A. Hooker of this Bureau. The 
manuscript is a study of the hfe history and habits of the North Amer- 
ican fever tick, tog-ether with notes on other species. The work upon 
which this bulletin is based was begun in July, 1905, after practically 
all of the directors of the southern experiment stations had brought 
to the attention of this Bureau the necessity of additional work on 
the important parasite which transmits Texas or splenetic fever of 
cattle. Prof. H. A. Morgan, director of the Tennessee Experiment 
Station, has given valuable advice during the progress of the work. 
The paper contains information of great value in the practical work 
of tick eradication. I therefore recommend that it be issued as 
Bulletin No. 72 of this Bureau. 

Respectfully, C. L. .Marlatt, 

Acting Chief of Bureau. 
Hon. James Wilsoa, 

Secretary of Agriculture. 



C()\T]-:\TS 



('()N(;kknin(; tiik north amkhican kkvkr tick. 

Puice. 
Inlroduclory 9 



occasioned by the calllc I ick 11 

The life history of ticks in general j 2 

The life history of the cattle tick j 3 

Period previous to oviposition 14 

Oviposition period J4 

Egg stage 15 

Incubation 17 

Relation of temperature to incubaiinn 19 

Effect of heat and cold on eggs 21 

Submergence of eggs in water 22 

Percentage of eggs hatching 23 

Larval or seed-tick stage 2o 

Nonparasitic period [ _ 23 

Effect of water on seed ticks 24 

Longevity of seed ticks 25 

Parasitic; period 27 

Development on liost 28 

Adidt stage 30 

Effect of continuous cold and lieat on engorge(l fenudes 31 

Effect of direct sunlight on adults 32 

Effect of submergence in water on engorged adult licks 3,2 

Dropping from host 33 

Locomotion " 33 

Host relations of the cattle t ick 34 

Relation between ration and tick infestat ion 35 

Enemies of ticks 3(; 

The practical application of llie information recorded in this bullelin 37 

X(rrK.S ON VARIOUS SPECIES OF Tl( KS FO I XD l\ IIIF: I \ FP K I > STA'J'KS. 

Classification and habits of ticks 40 

Key to families. sul)families, and .\ortli American genera of ticks 

(Ixodoidea) 40 

Family Argasidse 41 

Genus Argas 42 

Genus Ornithodoros _ 45 

Family Ixodidse 40 

Subfamily Rhipicephalina' 47 

Genus Rhipicephalus 47 

Genus Margaropus 49 

Genus Dermacentor 49 

Genus Haemapliysalis 52 

Subfamily Ixodinye 54 

Genus Ixodes ' 54 

Genus Amblyomnia 58 

Bibliography (;4 

Index 77 



ILLUSTRATIONS 



PLATES. 

Page. 

Pi,ATK T. Tlie North American fever tick. Fig. 1. — Eggs of the tick, Marga- 
ropus annulatus, deijosited under stable litter. Fig. 2. — Seed 

ticks of Margaropus bunched on grass and stake IG 

II. The North American fever tick. Fig. 1. — Steer used in experimental 

work. Fig. 2. — Arrangement for obtaining data on incubation .. . 16 

III. The North American fever tick and other species. Fig. 1. — Marga- 

ropus annulatus, male. Fig. 2. — Hsemaphysalis leporis-palustris, 
female. Fig. 3. — Stigmal plate of Margaropus annulatus, male. 
Fig. 4. — Mouth parts of Ixodes cookei. Fig. 5. — Stigmal plate 
of Rhipicephalus sp., male. Fig. G.— Stigmal plate of Ambhjomma 
maculatum, female 48 

IV. Stigmal plates of ticks. Fig. I.— Stigmal plates and anus of Dcrma- 

rentornitens,msde. Fig. 2.— Stigmal plate of same. Fig. 3.— Stig- 
mal plate of Amblyomma cajennense, male. Fig. 4. — Same, female. 
Fig. 5. — Stigmal plate of Dermacentor variabilis, female. Fig. 
G. — Stigmal plate of Dermacentor occidentalis 48 

TEXT FIGURES. 

Fig. 1. Margaropus microplus: Genital apparatus 14 

2. Graphic table for the separation of the families and genera of licks 41 

:'.. Rhl pi(( phnlus sp. : Capitulum of female 47 

4. Uliipiri jihiilus sp. : Coxae of male and female 48 

5. D( nitutxutor nitens: Capitulum of female 52 

6. Dermacentor nitens: Coxge of male and female 52 

7. Hsemaphysalis leporis-palustris: Capitulum and scutum of female 53 

8. Hxmaphysalis leporis-palustris: Coxse of male anrl female 53 

9. Amblyomma cajennense: Mouth parts of male 60 

10. Amblyomma cajennense: Coxse of male and female 61 

11. Amblyomma maculatum: Mouth parts of female 62 

12. Amblyomma maculatum: Coxiv of male and female 63 

13. Amblyomma maculatum: Scutum of female 63 

7 



INFORMATION CONCERNING THE NORTH AMERICAN FEVER TICK, 
WITH NOTES ON OTHER SPECIES. 



INTRODUCTORY. 

It is safe to state that no more important pi'ohleiii than the eradi- 
cation of the cattle tick (Margaropus '^ annulatus Say) confronts the 
farmers of any country. Not only the cattle-raising' industry, but 
the whole economic condition of a large section of coiuitry is affected. 
The tick, without unv but the most limited power of locomotion, 
and for all practical purposes dependent upon cattle for its (Existence 
and dissemination, presents a problem in eradication of a hopel'ul 
nature. Cattle are under the control of man. Therefore, the ])rot)- 
lem is quite different from that involved with other ])ests, like tlu^ 
boll weevil, which by flight spread over large areas of land. In the 
one case absolute eradication is possible and in the other it is out of 
the question. In fact the })()ssibility of the total extermination of 
the tick in this country is by no means visionary. It was foreseen 
originally, probably, by Dr. Cooper Curtice, who wrote as follows in 
1896: "I look most eagerly for the cleansing of even a certain portion 
of the infected territor}^ under the direct intention of man, for it 
opens the way to pushing the tick back to the S})anish Isles and 
Mexico, and hberating cattle from disease and pests and the farmer 
from untold money losses. Let your war cry l)e, ])eath to the 
ticks." '' 

In view of these facts it is evident that the most complete knowl- 
edge of the habits and life history of the tick is of the utmost impor- 
tance. All means of eradication must depend upon such knowledge, 
and improvements in present methods must depend upon additional 
information regarding the tick. Dr. Cooper Curtice, who will be 
quoted frequently, because he has been among the foremost in the 
study of the problem, has written as follows: ''To the scientist 
studying the tick to learn its life histor^^, habits, form, and anatomy, 

"Neumann has shown tliat the generic name Boophilus of Curtice must, in ol)e(li- 
ence to the zooloo;ical law of priority, fall as a synonym of ihc earlier naiiip, Marga- 
ropus of Karsch. 

^Journ. Comp. Med. and Vet. Archives, Vol. XVll, p. G.%. 



10 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

the fact that these animals are pests to the stockman throughout 
the greater part of the year is of very httle importance, while the 
latter cares little about such matters if he can only learn how to rid 
his cattle of them. Yet it is only by learning the life liistory that 
remedies to prevent them can be applied intelligently, and the fact 
that the knowledge attained is of practical value adds a double 
interest to their study." "^ 

In view of the evident importance of the work it is surprising that 
so little has been done in this country. In 1892, about a jenr pre- 
vious to the issuance of Smith and Kilborne's epoch-making bulletin 
demonstrating the tick transmission of fever, Dr. Cooper Curtice 
})ublished the first data regarding the life history of the cattle tick 
as Bulletin 24 of the Texas Agricultural Experiment Station. It 
was accompanied by excellent illustrations. The value of this work 
will be understood from the fact that it was of a pioneer character, 
and that all subsequent work has depended upon it. Nevertheless, 
it was of a preliminary nature and merely outlined matters that 
must eventually receive the most careful investigations. 

Prof. H. A. Morgan, principally in bulletins 51 and 56 of the Louis- 
iana Experiment Station, has added greatly to our knowledge of the 
cattle tick as well as other species. His work has such practical 
bearings that it has been the chief indication of the value of life- 
history studies in pointing out successful methods of eradication. 
Recently Messrs. Wilmon Newell and M. S. Dougherty, of the Louis- 
iana crop-pest commission, have published a valuable contribution 
which still further shows how every fact relating to the tick can be 
utilized in combating it. 

The above are the ijrincipai publications by American workers. 
There are many others which also contribute important facts. Among 
these are Connaway's, Schroeder and Cotton's, Ransom's, and others. 
In other countries excellent work has been done on related forms. In 
South Africa Prof. C. P. Lounsbury has macle scholarly studies of 
Afargaropus {Boopliilus) dexoloratus and many other species. In 
Argentina, Dr. F. Lahille has recently published the results of some of 
the most exhaustive work on ticks that has been done. These works, 
with others, are listed in the bibliography at the end of this bulletin. 

Notwithstanding the studies that have been conducted in this 
country, it must be stated that our knowledge of the tick is far short 
of what it shoidd be. There is a lack of knowledge of local variations, 
due to climatic influences, as well as such matters as dissemination. 
To supply this tleficiency, the Bureau of Entomology, in cooperation 
with the officers of several experiment stations, has undertaken a 



'I Tex. Agr. Kxp. Sla. Hul. 24, p. 238. 



LOSSES OCOASIONKD BY CATTLE TICK. 1 1 

careful study of the tick. Some of the results of practical bearino- 
are given in the following pages and others will be published from 
time to time. 

The writers desire to express their thanks to Prof. H. A. Morgan, 
director of the Tennessee Agricultural Experiment Station, for many 
most valuable and courteous suggestions in the course of this work. 
He has turned over to the wTiters many of his original notes and lias 
generously assisted in numerous other ways. 

LOSSES OCCASIONED BY THE CATTLE TICK. 

Undoubtedly the poi)ular idea of the damage caused by the cattle 
tick concerns itself with the actual death of cattle from the disease 
transmitted by the tick. Although this is a very important matter 
and would fully justify the most energetic attempts toward the 
eradication of the tick, it is really unimportant in comparison with 
the other losses. Mr. August Mayer, a practical' cattle breeder of 
Shreveport, La., and Dr. J. R. Mohler, of the Bureau of Animal 
Industry of tins Department, have made most careful, comprehensive 
estimates of the losses caused by ticks. The following summary is 
taken largely from their writings: 

1. Loss by death from disease in young animals and those removed 
from temporarily tick-free localities (as, for instance, in cities) to 
places where they become infested. The enormous loss under this 
heading will be understood when it is recalled that every bovine 
animal in the tick area must suffer an attack of fever if it becomes 
infested with ticks. In an instance that came to the attention of the 
writers, 39 out of 40 calves dropped in a city died of tick fever when 
removed to an infested pasture. 

2. Loss in weakened condition and stunted growth caused by the 
fever. 

3. Loss by gross tick infestation. At the present time (March, 
1907) hundreds of cattle in south Texas are dying from gross infesta- 
tion resulting from a mild winter. In extreme cases, Mr. Mayer 
estimates that as many as 200 pounds of blood may be withdrawn 
from the host during a single season. This makes a gain in weight 
impossible even in the best of pastures. Moreover, Prof. H. A. 
Morgan and other observers believe that gross infestation and the 
consequent general debility induce acute attack of fever even in 
animals ordinarily immune. 

4. The tick makes hazardous the importation of pure-bred cattle. 
This prevents the upbuilding of southern cattle and at the same time 
largely deprives the northern breeder of a market that he should have. 
Moreover, the inability of the southern breeder to exhibit his stock in 



12 NORTH AMKKIOAX FEVER TICK AND OTHER SPECIES. 

the north and of the northern breedei- to exhibit his in llie tick ai-ea is 
a handicap, the importance of which will be readily seen. 

5. The necessary restrictions in the shipping- of southej-n cattle 
also handicap the breeder and affect the price. 

6. The maintenance of the cjuarantine involves c-onsiderable annual 
expense for the protection of the cattle owners north of the line. 

7. Minor losses may be grouped as follows : (a) In Texas, especially, 
the tick induces the attack of the screw- worm fly (Chrysomyia macel- 
laria Fab.); (h) there seems to be, as pointed out by Mr. Mayer, a 
considerable interf^^rence with the fecundity of infested cows; (c) the 
railroads are put to the expense of disinfecting cars and maintaining 
separate pens and the stockman to the expense of dipping — items 
which react on the price that southern cattle bring. 

All the losses that have been mentioned total a])])roximately 
$100,()()0,0()() each year. At jn'esent the loss, as indicated by Doctor 
ISIohler, amounts annually to at least 10 per cent of the value of the 
cattle. The (|uality of tlie animals is the lowest and the loss is greatest 
in the regions wIkm'c the natural conditions without the tick should 
])roduce the iinest cattle ^\ith the least loss. But the damage may be 
better expressed by the stat(Mii(Mit that the tick makes profitable 
])roduction practically im])()ssil)le in the South. Any successful 
system of agriculture nnist rest uj)on a diversification of crops, and 
this, in turn, depends upon animal husbandry to maintain the fertility 
of the soil. Therefore, until the tick is (U'adicated oi' j)laced under 
control, a rational .system of agriculture in the infested area is out of 
the ({uestion, and that achievement would mean almost as much to 
the North as to the South. 

THE LIFE HISTORY OF TICKS IN GENERAL. 

The following genei'al statement I'egai-ding the life liistoiy of ticks 
is taken fi'om Salmon and Stiles:" 

Ticks arc temporary parasites, attacking inamiuals. birds, and reptiles. Tliey do 
not appear to be so strictly confined to ('(nlain hosts as do parasites in general. Still, 
this may be more of an apparent than a real rnle. Certain it is that, although a given 
tick may he found occasionally on aTiiinals which are very dissimilar (dog ticks have, 
for instance, been found on snakes), still the ^arious s^peeies show a decided predilec- 
tion for certain hosts. 

The parasites copidate during the i)eriod of parasitism '' and suck the blood from 
their hosts. The female grows to a large size and eventually drops to the ground and 



« Seventeenth Ann. Rej)t. Bureau of Animal Industry, U. S. Dept. Agric, p. 398. 

''This is not invariable. Arnhhjomnia aincrkanum scjmetimes copulates soon after 
the second molt, but f)efore it has gained a lio.st. It is likely that other sj)ecie.s also 
occasionally do so.— \V. D. H. and W. A. 11. 



LIFE HISTORY OF CATTLE TICK. 13 

lays numerous eggs, which are usually more (h- less clustered together. The larva 
upon hatchhag possesses three pairs of legs, the fourtli pair being added. during I he 
first molt. Either the hexapod or the ortnpod form may allack its host. 

From the foregoing it will he seen that the eatth^ tick, lik(> other- 
ticks, passes thi'ough tlie following stages: Egg, larva (six-legged 
form), nymph, and adult. 

The eggs are nearly ronnd, dark hrown in color, and (h^posited in 
largo masses, held together hy the gtmnny secretion with wliich the 
female coats each egg as it is (le])osited. The next stage, known as 
the seed tick, differs remarkably from the later stages in the fact that 
six instead of eight legs are present. The stigmata are located be- 
tween the second and third coxa>, just anterior to the third coxa-, 
and problematic indications are seen between the first and second 
coxa\ No distinct genital or anal opening can be seen in this stage. 
The anterior legs are much larger than the others. They are waved 
violently tlu-ough the air when the seed ticks are distiu'bed either l)y the 
approach of a host or in any other way. After some time th(> seed 
tick molts and the next, or n^nnphal stage, is provided with eight 
legs. The absence of the genital opening will differentiate this stage 
from the following one. Ticks in the nymphal stage are frequently 
referred to in the South as "yearling ticks." After a second molt 
the adult form is reached. Copulation then takes place, and after 
engorgement the female drops to the ground for the purjjose of depos- 
iting eggs. 

THE LIFE HISTORY OF THE CATTLE TICK. 

As })ointed out by Morgan the most important fact about the (lattle 
tick {Margaropus annulatus Say), from the stand})oint of ])ractical 
control, is that the time of development on the animal is always shorter 
than the total of the preoviposition, oviposition, and incubation 
periods. This gives the farmer an opportunity to free his cattle ajid 
pastures of ticks l)}^ the same process of rotation. As a foundation 
for the surest and most economical procedure an a{;curate knowledge 
of the variations of the periods in the life history of the tick under 
different conditions is absolutel}" essential. Our effort in this bidletin 
is in a measure to supply this information. The work has been prin- 
cipalh" to obtain data necessary in the pasture eradication and feed- 
lot systems of eradication. We have consequently studied the devel- 
opment of the tick both during its existence on the animal, by means 
of a steer procured for that purpose, and during its life, under various 
conditions, when not attached to the host. 



14 



NUKTH AMERICAN FEVER TICK AND OTHER SPECIES. 



PERIOD PREVIOUS TO OVIPOSITION. 



The cattle tick, like other species, passes tln'ough a distinct i)eri()cl 
between the time of dropping from the host and the beginning- of ovi- 
position. When the tick drops, the eggs are not ready to be depos- 
ited, but must pass from the ovary through the oviduct. (See fig. 1.) 
Thus there is a definite physiological basis for a period which has a 
very practical l)(>aring on ])lans of eradication that depend upon a 

knowledge of the exact time 
to be allowed in removing 
cattle fi'om one inclosure to 
another. Lahille has used 
the term "prootoquie" for 
this period, but we shall refer 
to it merely as the preovipo- 
sition period. 

As will be seen from Table 
I the preoviposition period 
ranges from 2 to 40 days, 
depending upon tempera- 
ture. In the summer it 
averages between o and 4 
days, and in winter over 20 
days. 

It might be supposed that 
the data in the table referred 
to show a preoviposition pe- 
riod longer than normal on 
„,,.,, . , ., . account of the removal of 

Ju;. ].-l.enital apparatus of Margaroimf: mtcroplus: a, 

Position of eggs at time of dropping of tick from host; the tlcks artificially. HoW- 

b. position of eggs when oviposition begins. Uighly ever, oulv ticks about tO drop 
magnifipa (redrawn from Lahille). ' ^ 

^\ere selected, and repeated 
tests with ticks actually dropped sIiowcmI that the method followed 
gives the natural preoviposition period. 




OVIPOSITION PERIOD. 



As will be seen from Table 1, the j)eriod occupied in oviposition 
ranges from 6 to 70 da^'s, depending upon the temperature. In the 
summer it averages 10 or 11 days, while in the winter it is two or 
three times as long. « 



LIFE HISTORY OF CATTLE TICK. 



15 



Table I. 



-Oviposition of Margaropus annulatus. July, 1905, to July, 1906, at 
Dallas, Tex. 



When collected. 



1905. 
July 21.... 

Aug. 3 

Sept. 18... 
Sept. 27... 
Oct. 12.... 

Nov. (■) 

Dec. f) 



1900. 

Feb. 6 

Feb. 21.... 
Mar. 23.... 

Apr. 6 

Apr. 20.... 

May 4 

May 22.... 

June 5 

Juie20 

July2 

July 13.... 



To'^al... 
Averssp- 



Preoviposition 
period. 



Days. 
3 



Days. 
2 
3 
3 
3 
4 

10 
21 



Days 
2.9 
3.5 
3.4 
3.0 
5.1 
11.9 
28.6 



Oviposition 
period. 



Days. 
13 
11 
18 
19 
44 
04 
70 



[^^ 



Days 
6 
7 
12 
19 
21 
21 
20 



Days. 
8.2 
9.0 
15.0 
19.0 
30.7 
39.9 
42.0 



23.4 
21.8 
15.6 
14.6 
12.6 
10.9 
11.7 
11.0 
10.4 



Period from drop- 
ping to end of ovi- 
position. 



Days. 
14 
13 
20 
21 
48 
79 



Days. 
8 
U 
14 
21 
21 
33 






Days 
11.0 
11.8 
17.0 
21.0 
32.0 
.50.8 
71.0 



.50.5 
47.8 
30.2 
25.7 
20.0 
21.4 
16.4 
13.4 
14.2 
13.9 
12.5 



Number of eggs per 
tick. 



Eggs. 
3,800 

2,228 



3, 875 
2,311 
2.689 
3,946 



2,134 
3,496 
2,437 
2,260 
3,412 
2,070 
3,180 
2,881 
2,407 
2,292 
2,397 



515 
1,460 



147 

10 

1,118 

1,281 

2,197 

1,701 

1,391 

152 

843 

1,135 

1,060 



Eggs. 
1,185 
1,971 



1,779 



2,009 
1,941 
1,058 
2,891 
2,251 
2,2.50 
1,802 
1.950 
1,837 
2.068 



32, 499 
1,911.7 



From Table I the following important, practical data are obtained: 

1. The preoviposition period ranges from about 3 days in summer 
to as many as 28 days in winter. 

2. The oviposition period ranges from between 8 and 9 days in 
summer to 42 days in winter. 

3. The total period from dropping to the end of oviposition ranges 
from 11 days in summer to 71 days in winter. 

It should be noted that Table I gives the total period from drop- 
ping to the end of oviposition based upon the weighted averages 
of the preoviposition and the oviposition periods. Therefore the 
maximum total period may be somewhat longer than indicated, as, 
for instance, in cases where either the preoviposition of the oviposi- 
tion for some reason are prolonged beyond the average. 

EGG STAGE. 



The eggs are generally elliptical, but vary in shape on account of 
pressure and drying. In color thej are at first honey-yellow, but 
soon change to a deep yellowish brown. They are shiny and 
smooth. The average size in a lot of 10, measured by a micrometer^ 
was 0.54 by 0.42 mm. About the middle of the incubation period 
in many species a whitish spot appears on the eggs and becomes more 
conspicuous as the time for hatching approaches. This spot is 



16 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

located toward one end and seems to be due to the excretion of the 
embryo. In Margaropus anmdatus it is very conspicuous and is a 
certain indication of viabihty. 

The act of oviposition is most interesting. This process was 
referred to by Dr. Cooper C^n^tice." An analogous operation in 
Ixodes ricinushns recently been carefully described and illustrated 
by \^aieler/' and was earlier noted by Lewis.'' One of our asso- 
ciates, Mr. R. A. Cushman, has observed the operation. The follow- 
ing description is l)ased upon his notes : 

When oviposition is about to take place the capitulum is bent 
downward toward the genital aperture. This exposes a delicate, 
viscid membrane betwecMi the capitulum and the scutum. The 
membrane becomes distended and is projected out over the capitu- 
lum in two round.Ml l<>b(^s. practically covering it. This process is 
repeated several tiim>s b<>fore the (^o-g is finally ejected, the mem- 
brane being extruded and ivti-acted alternately while the capitulum 
is lowered and rais(Ml. Finally tlu^ white, membranous ovipositor is 
exten.led, turning inside out. until it touches the distended mem- 
brane. T1h> capituhun is new completely hidden. As soon as the 
ovipositor and membrane have come in contact the former slowly 
recedes, heaving the (>gg adhering to and partially enveloped by the 
ineml)rane. The egg riMuains in this i)()sition for a varying length of 
time. Then the membrane is withdrawn, rolling the egg along for a 
short distance on the dorsal surface of the capitulum. At the same 
time the ca])itulum is raised. Then the ])rocesses of distention and 
contraction of the membr:uie and lowering and raising of the capitu- 
lum aiv repeated s(>v(n-al times, the egg Ijeing finally completely 
coated by the viscid substance from the membrane and being finally 
pushed i)ack and d(>i)osited on \]\v anterior edge of the scutum. 
p:ach egg is laid in this manner, the tick backing slowly away and 
leavingMlie mass of eggs in front of her. The actual time consumed 
by the tick in laying a single egg is about 80 seconds, w^hile the 
removal of the egg and the resting period consume from one to sev- 
eral minutes, a much longer resting i)eriod being taken at intervals 
between lots of from 10 to 50 eggs. It has been impossible for us 
wdth the means at our command to demonstrate the ''paired, race- 
mose glands" of the membranous sac referred to by Curtice. As far 
as we have been able to s(>e, tho substance with which the eggs are 
coated is secreted from numerous minute glands scattered over the 
surface. 



"Tex. Agr. Exp. Sta., Bui. 24, p. 242. 
?>.Tourn. Agric. Sci., 1906, p. 405. 
cProc. Rov. Mici-os. Soc, 1892. 



3logy, U. S^ Dept. of Agriculture. 



Plate I. 




Jul. 72, Bureau of Entomology, U, S, Dept. of Agncultun 



Plate II. 




LIFE HISTORY OF CATTLE TICK. 17 

In the manner described a mass of eggs grows steadily in front of 
the tick, while its body becomes correspondingly smaller as the 
process proceeds. The gummy secretion holds the eggs together so 
that the mass looks not unlike a large accunudation of minute brown 
beads. 

The number of eggs deposited varies greatly with the size of the 
female. The highest number recorded in our experiments was 3,806." 
The average of 189 ticks under various conditions was 1911.7, and 
this probably very closely approximates the average under natural 
conditions. The daily average, of course, varies also. The maxi- 
mum is generally reached from 7 to 9 days after deposition begins. 
The highest number for any 24-liour period was 826. The average 
for 20 ticks was 144. 

INCUBATION. 

Most important means of control of the cattle tick depend upon 
taking advantage of the fact that eggs remain on the ground for a 
considerable time before hatching. Provided there are no seed ticks 
present, it is perfectly safe to allow cattle in areas in which ticks may 
be dropped from them, as, for instance, in fields under cultivation 
for one crop season, if the animals are removed before hatching takes 
place. It will be seen that this has an important bearing on the 
process of relieving cattle of ticks by placing them for limited periods 
in different tick-free inclosures. Accordingly we have made an 
especial effort to obtain data regarding the period occupied in incu- 
bation under different conditions and in different seasons. 

In 1905 a number of experiments to determine the length of the 
incubation period were conducted, the eggs being placed in paper 
pill boxes. Subsequent work showed that this arrangement gives 
more rapid development of the embryo than takes place under normal 
conditions. Especially is the period shortened when an abundance 
<jf moisture is furnished. In these experiments in July and August 
the eggs hatched in from 17 to 21 days, and during September in 
from 25 to 44 days. These results are of value only in showing how 
the incubation period may be shortened under extreme conditions, 
which must rarely or never occur in nature. 

No eggs deposited in October, 1905, hatched before April 10, 1906, 
a period of over 170 days. 

a The greatest number found by H. A. Morgan was 3,198 (La. Agr. Exp. Sta., Bui. 
51, p. 242), but Newell and Dougherty record 4,124 in one instance (La. Crop Pest 
Comm., Circ. 10, p. 23). 

5795— No. 72—07 2 



18 NORTH AMERICAN FKVER TICK AND OTHER SPECIES. 

In 1906 a more extended .series of experiments was instituted. In 
the first series, consisting of 59 lots deposited from February 20 to 
September 21, eggs were kept in pill boxes. As has been pointed out, 
this method accelerates incubation further than natural conditions 
are likely to. The following is a summary of the results of these 
experiments : 

Eggs deposited in March hatched in 74 day.s. 

Eggs deposited in April hatched in 43 to 53 days. 

Eggs deposited in May hatched in 24 to 33 days. 

Eggs deposited in June hatcned in 20 to 22 days. 

Eggs deposited in July hatched in 19 to 22 days. 

Eggs deposited in August hatched in 19 to 21 days. 

Eggs deposited in September hatched in 23 to 154 days. 

Eggs deposited October 1-7 hatched in 135 to 139 days. 

Eggs deposited October 15 and later have not hatched to date (Feliniary 20, 1907). 

In the second series of experiments relating to incubation conducted 
in 1906 a special effort was made to provide conditions that would 
give approximately the same period that must occur under normal 
ccmditions. (wSee Table II.) To do so some eggs were placed on soil 
in o])en-bottom glass tubes in the open air (see PI. II, fig. 2) and shaded 
for a portion of the day. Others placed under the same conditions 
were exposed to the sun at all times. In the third series the eggs 
were located in an outdoor thermometer shelter where they were pro- 
tected from sun and rain. The separate lots consisted of several 
hundred eggs. They were deposited by different femal(»s in the 
laboratory and placed together for the observations under considera- 
tion. In this way a large number of eggs deposited within the same 
24-hour period was obtained. It is supposed that these conditions 
approximate closely to those which surround the great majority of 
eggs ill pastures. The following is a simimary of the data contained 
in the table: 

Eggs deposited in April hatched in from 39 to 54 days. 
Eggs deposited in May hatched in from 27 to 33 days. 
Eggs deposited in June halcheil in from 21 to 28 days. 
Eggs deposit (>d in July hatched in from 22 to 2(5 days. 
Eggs depositt'd in August liatchcd in from 23 to 32 days. 

In September eggs deposited prior to the 18th hatched in from 23 
to 76 days. Eggs deposited after September 18 have not hatched up 
to February 20, with but two exceptions. These were eggs deposited 
October 3 and October 7, wliicli began to hatch on February 18. 
These eggs were placed in such a way that they were more exposed to 
the sun than other lots that have not hatched, deposited before and 
since the dates mentioned. 



LIFE HISTORY OF CATTLE TICK. 



19 



Table II. — Period of incubation of Margaropua nnnulnlii)t at Dnllns, Tex., 1906. under 
various conditions. 



iCggs (Ipposited. 



Apr. 13. 
Apr. 14. 
.\pr. 1.2. 
-\pr. 2/ . 
May 13. 
May l.T . 
May 28. 
May 3iJ. 
May 31 . 
June 9.. 
.June li». 
June 1 1 . 
June 21 . 
June 22. 
June 24 . 
June 24 . 
June 30. 
July 1 . . 
July 20. 
July 20. 
July 22. 
Julv 31 . 
July 31 . 
July 31. 
Aug. 1 1 . 
Aug. 12. 



Ilatdiing. 



Minimum | 
iiu-ubation i{ Kggs dcii 
period. ; 



June o . . 
May 30. 
May 31.. 
June 4.. 
June 14. 
June 12. 
J une 23 . 
June 26. 
June 27. 
Julys.. 
July 5.. 
July 7.. 
July IK., 
July 14.. 
July 16.. 
July 16.. 
July 20. 
July 22.. 
Aug. 10. 
Aug. 14. 
Aug. 14. 
Aug. 22. 
Aug. 22. 
Aug. 23. 
Sept. 4. . 
Sept. 3.- 



Days. 



a 29 
127 
128 
"28 
127 
"26 
127 
128 
''23 
123 
''23 
''21 
122 
''22 
126 
C24 
''23 
C23 
124 
''25 



Aug. 
Aug. 
Aug. 
Aug. 
Aug. 
Aug. 
Aug. 
Aug. 
Aug. 
Aug. 
Aug. 
Aug. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 
Sept. 



Sept. 8. . 
Sept. 6.. 
Sept. 9.. 
Sept. 18. 
Sept. 10. 
Sept. 11. 
Sept. 13. 
Sept. 14. 
Sept. 10. 
Sept. 20. 
Sept. 21. 
Sept. 23. 
Sept. 23. 
Sept. 24. 
Sept. 27. 
Sept. 26. 
Oct. 7... 
Oct. 10.. 
Oct. 15 . . 
Oct. 24.. 
Oct. 26.. 
Nov. 5.. 
Dec. 2... 
Dec. 2... 
Feb. 20.. 



Minimum 

incubation 

period. 



Day.'i 



i2S 
''24 
''23 
132 
''23 
f24 
123 
C24 
''24 
''24 
C25 
127 
C23 
6 24 
127 
-^25 
C29 
ft 32 
137 
c41 
''43 
153 
''76 
i76 

''148 



1 The eggs were placed in open-bottom test tubes in sand in the open air. They were protected from 
the direct rays of the sun at all times by a cheese-cloth screen, fp to 11 a. m. they were 
within the shade of the house. 

'' In open-bottom test tubes in soil exposed to sun at all times. 

c In open pill boxes outdoors protected from sun and rain at all times. 

RELATION OF TEMPERATITRE TO INCUBATION. 



In th(^ series of experinieuts just referred tt) accurate data on tem- 
perature were obtained. Standard maximum and minimum ther- 
mometers kept in an instrument shelter were used. In Table III 
the records of temperatures are given together with the average 
incubation periods for the various lots of eggs imder observation 
during different months. The data show that there is an intimate 
relation between temperature and the period of incubation. The 
shortest average minimum incubation period (23.4 days) occurred 
when the average temperature was liighest (80.2° F.). The longest 
average minimum incubation period (137 days) occurred with the 
lowest average mean temperature (53.2° F.)., Between these ex- 
tremes there is a graduated correspondence between temjierature and 
incubation. 



20 



NORTH AMKRI(;AN FEVER TICK AND OTHER SPECIES. 



Table III. — Relation of tcniperaturr to period of inriibntion in Margaropiis annulatu^ 
at Dallas. Te.r.. 1905-1906. 



Month <lf])osi1f 



1!K).-, 
sq't 

v.m 

M^y.".v.' ■;;;;;;; 

June 

July 

Aug 

Sept 

Oct 



No. 

of 

lots. 



Incubation 
period. 



Maxi- Mini- Aver 
mum. mum. age.' 



Days 
4,3 



Incubation temperature. 



Total eflfeetive. 



Maxi- Mini- \ver- 
mum. mum. aee. 



Days. °F. 
49.5 62.6 



46.7 
29 
24.8 
23.4 
24.9 
46.2 
137 



79.7 
80.5 
80.5 
79.7 
79.4 
53.4 



Maxi- 
mum. 



°F. 
62.4 



69.7 


70.8 


73.5 


77.6 


79.8 


80.2 


V9.8 


80.2 


78.9 


77.8 


54.6 


70.7 


53 


53.2 



°F. 
, 0S3 



1,103.2 
1,046.6 
%2. 1 
1,169.3 
1,907.9 
1,600.6 



Mini- Aver- 
mum. I age. 



1,139.1 
981.6 
782.7 
824.3 
826.7 
837.6 

1,510.8 



1,286.4 

1,023 
920.7 
866.5 
904.4 

1,127 

1,5.55.7 



" WcightPd. 

It will he noted that the total effective temperature necessary to 
cause eggs to hatch varies from 866.5° to 1,555.7°. It would be of the 
greatest practical importance to formulate a rule w^hich, on the basis of 
effective^ temperatures, would show the time when eggs will hatch at 
different seasons of the year. It is probable that the present data are 
insufficient for this purpose, and special efforts will l)e made to add 
to them. However, with the data at pivsent available the follow ing 
tentative law may be ])roposed : " 

Whe7i the average daily mean temperature ranges less than dSJ°, at 
least 1,510.8 degrees of effective temperature must accumulate hefore 
hatching tvill take place. When the mean daily temperature averages 
from 61 .4° to 77.8°, from 8Jfi.5 to 1 ,139.1 degrees of effective temperature 
will he required for hatching. \yhen the mean daily temperature aver- 
ages higher than 80°, between 782.7 and 82If..S degrees of effective tem- 
perature must he accumulated hefore hatching will take place. 

It is not our purpose to advise an attempt at the practical appli- 
cation of this rule at the present time, but it is supposed that such 
])ractical application can be ultimately made. To do so it wamld 
only be necessar}" for the stockman to have a set of self-recording 
maximum and minimum thermometers, such as can be purchased 
for $3. Or the data might be obtained from the nearest Weather 
Bureau station. By either of these means the average daily tempera- 
ture could be* easily obtained. By summing up the daily effective 
temperatures — that is, the number of degrees above 43 — the stock- 
man, by reference to the minimum amount of accumulated effective 
temperature necessary for hatching with various average daily tem- 
peratures, could determine at least within certain limits what time 

« This rule, of course, may not apply in rare cases in which eggs are deposited where 
they will be suliject to artificial heat rather than to weather temperature, as in manure 
piles. It will apply, however, to the great majority of pasture conditions throughout 
the infested area. 



LIFE HISTORY OF CATTI.K TICK. 21 

it would be necessary to remove cattle from pastujvs in which tick 
eggs might be to avoid danger of infestation by seed ticks. Possibly 
a more feasible application would be the collection of the necessary 
data from many localities by the State entomologists and the ])ubli- 
cation of predictions based upon them from time to time. 

A seasonal arrangement of our data for 1905 and 1906 shows the 
following : 

Eggs deposited in June, July, August, and up to September 15 
reciuire from 824.3 to cS40.5 degrees of accumulated effective tem- 
perature for hatching. 

Eggs deposited from September 16-30, in October, and in later fall 
and winter months, require an accumulated effective temperatm-e of 
from 837.6 to 1,510.8 degrees. 

Eggs deposited in April and May require from 9S1.6 to 1,139.1 
degrees. 

Naturally an arrangement by months as above must be defective, 
since no two seasons are exactly alike. The only accurate method 
must be based upon a knowledge of thf^ temperatures that arc accu- 
mulating in any particular season. 

EFFEC'l' OF HEAT AND COLD OX E(i(JS. 

In experiments to detennine the effects of heat iq)on eggs a con- 
tinuous temperature of 100° was maintained by means of an incu- 
bator. The period of application of heat was 15 days. In one series 
no moisture was provided, and in this case no hatching took place. 
In another series in which abundant moisture was furnished, hatch- 
ing took place and the incubation period was reduced to 15 days. 
It will thus be seen that a moist atmosphere is essential to the hatch- 
ing of eggs under a constant high temperature. 

In experiments relating to the effects of low temperatures on eggs, 
by means of a refrigerator a mean temperature of about 45° was 
maintained, with a minimum of 32° and a maximum of 65°. The 
eggs were kept in pill boxes with gauze tops to allow free circula- 
tion, except during the period of refrigeration, when the ordinary 
covered pill boxes were used. The eggs from about 20 engorged ticks, 
collected on July 26, were placed in the refrigerator on August 4 
and remained for 30 days. Hatching began on September 23, and 
about 60 per cent of the eggs were viable. 

In the case just referred to, the normal period of incubation w^as 
increased, as the result of refrigeration, by 8 days. In a long series 
of similar experiments, however, in which the period of refrigera- 
tion ranged from 1 to 21 days, the period of incubation was not 
appreciably lengthened and the normal percentage of hatching took 
place. 



2:2 NORTH AMEKK'AN FEVEK TICK AND OTHER SPECIES. 

In a series of experiments with alternate cold and normal tempera- 
tures the eggs were kept in a refrigerator exposed to a mean tem- 
perature of about 45° F. from 8.30 p. m. to 8.30 a. m. During, the 
day the eggs were removed from the refrigerator and remained at 
the temperature of the air. After six consecutive nights of exposure 
in this way from 5 to 10 per cent of the eggs hatched. It is probable 
that the necessary manipulation in these experiments interfered 
with the viability of the eggs and that normalh^ a considerably greater 
percentage would hatch under the same conditions. 

Sl'BMERGENCE OE EGGS IN WATER. 

Sixteen different lots of eggs were used to determine the effect of 
submergence in water. One-half were submerged for from 10 to 24 
days, and most of the eggs hatched. In another lot submerged 
for 25 days 33 per cent hatched. The experiments were per- 
formed from June to September and the incubation period under 
water was not a[)preciably different from the normal at the cUfferent 
periods at which submergence took place. In all these experiments 
complete submergence was secured by means of a screen obstruc- 
tion below the surface of the water. Our results agree with those 
recently ]nd)lished by Messrs. Newell and Dougherty and with unpub- 
lished data obtained by Prof. H. A. Morgan, who suggested our 
experiments. 

The practical im])ortance of these experiments is to show that the 
flooding of ])astures would have no effect whatever on the viability 
of tick eggs on the ground. Not only would the great majority 
hatch, but the time of hatching would not be materially different 
from that in case no water whatever were present. As a matter of 
fact the flooding under some conditions, as, for instance, during a 
drought, might hasten incubation. 

These data, taken in c(mnection with (hita mentioned elsewhere, 
showing the remarkable resistance of seed ticks to water, indicate 
clearly the reason why pastures overflowed for considerable periods 
have repeatedly been found to furnish tick infestation. 

A series of eggs varying from those recently deposited to others 
about to hatch were submerged in tube-form vials. To keep the 
eggs submerged absorbent cotton was pressed down into the water. 
None of the eggs hatched, and w(» suppose this was due to insuffi- 
cient aeration. 

Several lots of eggs that were kept in pill boxes until they had 
become thoroughly dried were placed on water in Stenter dishes to 
determine if hatching would follow. After submergence for a short 
period the eggs in large part filled out and appeared viable, but did 
not hatch. 



LIFE HISTORY OF CATTLE TICK. 28 

PERCENTAGE OF EGGS HATCHING. 

To determine the exact percentage of eggs liatching is a rather 
(Hfficidt matter for the reason that manipuhition in counting inter- 
feres greatly with their viabiUty. In June, July, and August, 1906, 
a number of observations were made showing the percentage of 
viable eggs in different lots fi-om oO to 90. In these cases the eggs 
were counted, and it is likeh' tiiat the percentage hatching under 
natural conditions is considerably higher than that indicated. The 
examination of masses of eggshells where hatching has taken ])lace 
normally reveals few unhatched. eggs. 

Messrs. Newell and Doughert}^" have recorded a percentage of 
hatching during the months of April, May, June, July, August, and 
September of from 61.6 to 92. These experiments were performed 
at Baton Rouge, La., under conditions which approached the natural 
ones verv closely, although the eggs were counted as in our experiments. 

LARVAL OR SEED-TICK STAGE. 

The larval tick is a minute 6-legged creature without distinct genital 
opening, and with indistinct stigmata between the second and third 
pairs of coxa3. As Salmon and Stiles state, larval ticks frequently show 
indications, at least, of stigmata between the first and second pairs 
of coxae, and behind the posterior coxse in addition to those between 
the second and third pairs. Only the pair first mentioned seem to 
be functional. The color of the larva at first is whitish but soon 
becomes dark brownish. 

NONPARASITIC PERIOD. 

For a few hours the larva remains about the shell from which it has 
just emerged, but later makes its way upward on the first blade of 
grass, stick, post, or other support that presents itself. Professor 
Morgan informs us that he has seen seed ticks on the tips of sugar 
cane about 8 feet from the ground. By placing a pole in the vicinity 
of millions of seed ticks we have observed them to reach a height of 
about 6 feet in a surprisingly short time ; but the tendency is strongly 
to remain not more than about 4 feet from the ground. In the absence 
of some vertical object the seed ticks do not seem to scatter to any 
great extent, but collect on the highest immediate point, even if it 
is only a small clod or stone. 

On whatever support the young ticks happen to be, they collect 
in masses often nearly an inch in diameter. (See PI. I, fig. 2.) Here 
they remain for weeks or months awaiting a host. The front legs, 
wnich combine the functions of antennae and legs in the insects 

oLa. Crop Pest Coinm./circ. 10, p. 24. 



24 



NORTH AMEKTOAN FEVER TICK AND OTHER SPECIES. 



proper, nve waved through tlie air more or less constantly, and vio- 
lently when a moving object approaches. The ticks attach them- 
selves to any animate or inanimate object which touches them. Only 
those that happen to attach themselves to cattle (and rarely a few 
other animals) ever develop; the others either die or, dropping off, 
become widely scattered. 

Heavy rains wash the seed ticks to the ground, and it is possible 
that violent winds may also serve to disseminate them. In our 
experiments the}' have thus been spread to a distance of 5 or 6 feet. 

It is noticeable that the seed ticks shun direct sunlight. We have 
repeatedly seen bunches move halfway around the support with the 
shade. In the morning they would be on the west and at night on 
the east side. 

EFFECT OF WATER OX SEED TICKS. 



Interesting data having a bearing on the dissemination of the cat- 
tle tick through the agency of water courses have been obtained. It 
has been not(Ml that heavy rains wash seed ticks from their supports 
to iho ground. In a considerable number of experiments seed ticks 
were found to endure submergence varying in different lots from 
10 days to 157 days. The latter record was obtained in an experi- 
ment in which seed ticks were first placed in water in a Petri dish 
and a few days later removed to a tube with earth on the bottom. 
The details of these experiments are given in Table IV. 

It is doubtless true that dissemination by water courses is not 
quite as important as these results would indicate. Of course it is 
possible that seed ticks may be carried many miles and deposited 
on grass or bushes, from wliich they may reach cattle. The enor- 
mous scattering of ticks so submerged in water, as would be the case 
in floods, would undoubtedly greatly reduce the chances of infesta- 
tion in pastures. 

Table IV. — Effect of water on seed ticks. 




Eight inches of water in tul). 

In Petri dish; Oct. 9 removed to tube with dirt on bot- 
tom. 
Alive on Nov. 25, and may have Uved longer. 
In tub with sand and vegetable matter. 
In Petri dish on porch, with algae in water. 
In Petri dish on laboratory desk. 
Submerged as eggs. 
Submerged as eggs. Ciilicid larva may have interfered. 



Some years ago Professor Morgan obtained interesting results on 
the effect of low temperature on seed ticks. In brief, he found a 
temperature of 15° or 16° F. for a short period did not kill many seed 



LIFE HTSTOKY OK OATTLK TICK. 25 

ticks, but that such a temperature continued for 24 liours resulted in 
the death of practically all. Our experiments have been of two 
classes — (1) in water and (2) without water. In both cases the seed 
ticks were subjected to a temperature somewhat less than 32° F. In 
the case where water was supplied solid ice was <)])tained. Both in 
and out of water the seed ticks survived a freezin^: temperature of one 
hour's duration. althou*i,h there was ]>erceptible mortality among 
them. 

I.ONCJEVrPY OK SEED TICKS. 

The time that seed ticks may survive without a host is a most 
important matter in plans for control. Our experiments on this 
point have been of two kinds— (1 ) with seed ticks from eggs deposited 
by females placed on the ground in favorable circumstances, thus 
giving absolutely natural conditions, and (2) with seed ticks in glass 
tubes, where they could be observed more closely. 

In the first .series, which w^as instituted at the suggestion of Pro- 
fessor Morgan, several hundred engorged ticks wwo |)lace(l on tl;e 
ground at regular intervals. The only inclosure was a cylinder of 
2-inch-mesh wire screen about 4 feet in diaineter to pr-event (hsturb- 
ance. The seed ticks from females placed under tiiese conchtions 
accumulated in enormous numbers on the blades of grass or stakes 
provided for the purpose (see PI. I, fig. 2). Table V gives the num- 
ber of days the seed ticks survived in these experiments, together 
with other data. It wall be seen that the shortest period w^as 49 
days and the longest 159+ days. However, the important period 
is from the time of dropping of the adults to the death of the resulting 
seed ticks, since the farmer must always suppose in rotating his cat- 
tle that adults were dropped on the day of removal from the pasture 
which it is desired to clean. This period ranged from 91 to 175 + 
days. It will be noted that there is considerable variation in the 
period of survival, even at the same season of the year. This seems 
to depend upon two factors, namely, the number of seed ticks in the 
bunches and the amount of rainfall. The larger masses survive 
longer, perhaps because the moisture is better retained, and heavy 
rains scatter and reduce the masses. It is probable, on account of 
the very large numbers of seed ticks in our experiments, that the 
periods given in the tables are somewhat longer than normally occur 
in the field. How^ever, such excessive numbers do sometimes occur 
in nature. For instance, wdien an animal dies of gross infestation 
thousands of ticks deposit in a very restricted area. Mr. J. D. 
^litchell has seen cases of this kind in which the bunches of seed ticks 
were fully as numerous as in our experiments. 



t>(; 



NUKTH AMERICAN FEVER TICK AND OTHER SPECIES. 



Table V. — Longevity of seed ticks of Margaropus annulatus Say, at Dallas, Tex., 1906. 



When dropped. 


Period from dropping 
to oompletion of ovi- 
position. 


When 
hatched. 


Seed ticks 
all dead. 


Maximum 
period 

from drop- 
ping to 

death of all 

seed ticks. 


Period 

of life of 

seed 

ticks. 




Minimum. 


Maximimi. 


June 20 


Days. 

12 
12 
14 

S 


Days. 

18 
18 
21 
19 
10 
16 


Aug. 12... 
Aug. 15... 

Julys 

Aug. 14... 
Sept. 1.... 
Sept. 14... 


Oct. 8 

Oct. 21. . . . 
Oct. 20. . . . 
Oct. 15.... 
Oct. 20. . . . 


Days. 
110 
120 
154 
80 
91 
17.")+ 


Days. 


June 23 


a 67 


May 19 


107 






Do.. 

Aug. (i 


49 

1.59+ 









a In this experiment the seed ticks wore accidentally disturbed. They would have lived some days 
beyond the period indicated. 
i Some alive Feb. 20. 1907. 

Ill the second series t)f experiments to tleterniiiie the longevity of 
seed ticks the eggs were placed in glass tubes with open bottoms. 
As will be seen from the footnotes to Table VI certain seed ticks in 
tubes were shaded at all times, others in tubes were exposed con- 
stantly to the sun. while the remainder were placed in pill boxes 
protected from sun and ruin. It is supposed that these diverse^ con- 
ditions give an av(M-age lengtii of survival that approaches closeh" to 
that occurring under natural conditions. The detailed results follow: 

Tahm-; YI. — Longirily of .sc(d ticks of Margaropus armnlalus Say, Dallas, Tcr., I'KiG. 



Eggs deposited.' 


Hatching." 


: Period Period 

•^pod tipL.j<lp.i<i ' ^""o™ fromdep- 
Secd ticks dead. hatching ositiou 

to death, to death. 


Condi- 
tions. 


Apr. 13 




\ug 28 


Days. 

84 

83 


Days. 
138 
124 
1.30 


(6) 


May 30 




Acr. 15 


May 31 


Aug. 22 


(d) 








85 , 124 

98 i 131 

115 144 




Mav 13 


June 14 


Sept 20 


()j) 


May 15 


June 1'' 


Oct 5 


May 2S 


June 23 


do 


104 


131 


(b) 








{b) 


May 31 






43 
83 


110 








(fc) 


June 10 






(b) 


June 11 


July 7 




28 
45 
21 
82 
(i8 
00 
71 


104 
56 
08 
44 

105 
89 
88 
93 


(h) 


June 21 

J une 22 


July-lS 

July 14 


Aug. 15 

Aug. 28 


W 


Jun(> 24 


July 10 


Aug. 6 


(b) 


Do 


do 

July 20 : 

July 22 

Aug. 10 

Aug. 14 










( '") 


July 1 


do 

Oct. •'0 


(6) 


.July ''0 




"Do 




(b) 


July 22... 








(d) 


July 31 


Aug. 22 

do 


P'eb. 18 


180 


203 


(e) 


Do 




((/) 


Do 


Aug. 23 




153 


177 


[b) 









open air. They were protected from 
Up to 11 o'clock a. m. they were also 



a For details regarding oviposition of these ticks see Tables I and II 

b The seed ticks were in open-bottom test tubes in sand in the o 
the dlrpft rays of the sun at all times by a cheese-cloth screen " 
shaded by the house. 

c In open-bottom test tubes in soil exposed to sun at all times. 

d In open paper pill boxes outdoors, protected from sun and rain at all times. 

Considerable numbers of seed ticks in all lots hatching after August 
23, 1906, are still ahve (February 20, 1907). 



LiFE HISTOKY OF CATTLK TICK. 27 

PARASITIC PERIOD. 

The data already given regarding- the periods of preoviposition, 
incubation, and survival of seed titks have an important bearing on 
the time required to free pastures or other inclosures from ticks i)ro- 
vided the cattle are removed. The data given under the present head- 
ing, on the other hand, show the time required at different seasons 
to free cattle of ticks by placing them in inclosures from which the ticks 
have been eliminated either hy systematic starvation or hy the use of 
naturally tick-free areas, as, for instance, fields that have been in culti- 
vation for one crop season. 

In this work we have utilized a grade Durham steer, 17 months 
old at the beginning of the experiments. (See PI. II, fig. 1.) By 
means of kerosene emulsion he was carefully cleaned of the thou- 
sands of ticks infesting him when obtained. Thereafter he was thor- 
oughly washed to remove traces of the insecticide and hundreds of 
seed ticks were applied. Under proper precautions to avoid the steer's 
accidental infestation, these ticks were allowed to reach maturity. 
^\iter the ticks of each infestation became adult the steer was thoroughly 
cleaned and placed in another inclosure, which in each case had been 
carefully disinfected by means of spra3^s. This process has now been 
repeated until ten infestations have been reared covering the period 
between August, 1905, and March, 1907. The details are given in 
Table VII. In rotation systems the minimum developmental periotl 
is the most important, because the cattle must be removed l)efore 
the earliest developed ticks have had offspring to reinfest them. 
Therefore special reference is made in the table to the shortest periods 
found, although the longest and the averag(^ are both given. 

The following deductions may be made from this table : 

1. The period from attachment to dropping ranges from 21 to 58 
days. It should be noted that in the longest periods the limit was 
reached by only one or two belated ticks, the majority approaching 
the average. 

2. The average period ranges from 26.5 to 43 days. 

3. The average parasitic period is normally souk^ tlays longer in 
winter than in summer. But warm winter weather, as happened in 
infestation No. 9, may reduce the period even below the average for 
the summer. 

4. The slowest developing ticks of one infestation may occupy from 
10 days (in the summer) to 32 days (in the winter) longer than the 
most rapidly developing ones. The rapidity of development of the 
ticks of the same infestation depends somewhat upon their location. 
Those on the portions of the body where the blood supply is most 
abundant develop most quickly. In general it seems that heavy infes- 
tations tlevelop a little more quickly than light ones. This may be 



28 



NORTH AMERICAN FKYER TICK AND OTHER SPECIE^ 



due to the fact that, in hght infestations with widely scattered ticks, 
fertihzation is less likely to take place than in other cases where the 
males may fuid the females more readily. We have been unable to 
determine that unfertilized females occupy longer in development 
than those that are fertilized, but our impression is that they do. 

The table also shows that the principal variation in the time of 
development of the ticks of the same infestation takes place not in 
tho larval or nvmphal but in the a(hdt stage. 

Tahlk \ll.— I>< rflopiiinU of .\furyaropii.s tnuuilaltis on star at Dallas, Tex. 



Ap- 




ph- 
ca- 
tion 


When 
appliod. 


No. 




1 


1905. 
Aug. If) 
Sept. 27 
Nov. 11 


4 

(i 

8 
!) 
10 


1900. 
Jan. It) 

Aug. 2 
Sept. 5 
Oct. b 
Nov. 29 


n 


1907. 
Jan. 1 



Mini- 

inum Second 
larval I molt, 
stage. I 



Aug. 28 
Oct. 4 
Nov. 22 



Jan. 25 
May 31 



Days. 
12 



\ug.lO-ll 
Sept. 12 
Oct. 14 
Dec. 8 



Sept. 2 
Oct. 12 
Nov. 28 



Feb. :i 

June I) 

July 10 

Aug. 18 

Sept. 22 

Oct. 21 

Dw. 14 



Adults dropped. 
Mini- 


Mini- 








nym- 

phal 1 First, 
stage. 


Last. 


pe- 
riod 
adult 

stage. 


Day.^. 




Days. 


5 Sept. 15 


Sept. 27a 


13 


8 Oct. 21 


Nov. 8 


9 


li Pee. S 


Jan. 9 


10 


ii Fell. 17 


Mar. 8 


14 j 


C. June 14 


.lune ;« 


8| 


Inly 24 


Aug. 5 


8 


7-8 Aug. 25 


Sept. 11 


7 


10 1 Sept. 2(i 


Ot^. U 


4 


7 Oct. :«) 


Nov. 12 


9 


11 Dec. 22 


Jan. 1 


8 


ti Jan. 24 


Kel). -.i 


10 



N^eTi l^"'od from 
"^ I attaciiment to 
,ov- dropping. 



, 1 drop- Maxi-, Mini- j Aver- 
ped. mum.:inum. age. 



Days. 
3() 
.■53 
43 



41.5 

31 

31 

31.5 

2(). 5 

:«. 5 

28 





Days. 


Days. 


20 


42 


30 


10 


42 


24 


13 


.'59 


27 


11 


51 


:g 


9 


;S9 


23 




37 


25 


14 


40 


23 


23() 


31 


21 


2(H) 


.37 


24 


55 


.33 


23 


35 


33 


23 



DEVELOPMENI" ON HOST. 



When the larval tick.s (hid t hciiis("lv(>s on the host they rapidly dis- 
appear in the liair and jittach themselves to the skin. They are 
principally found on such ])arts as tlie legs, bell}', and dewlap that 
come in contact with the bunches on the grass, but maybe found on 
an}' part of the host. In cases of severe infestation they practically 
cover the entire surface of the body, even the eyelids being infested. 

In from 7 to 12 days the larval ticks molt and enter the nvmphal 
stage, in which they have eight instead of six legs. The nymphal 
stage is further distinguished from the larval stage by the presence of 
a pair of large stigmata quite in c<mtrast to the riuHmentarv organs 
of respiration of the larva. 

The second molt (from the nymphal to the adult stage) occurs in 
from five to ten days after the first. The nymph can be distinguished 
from the adult, which it resembles very closely, l)y the absence of any 
genital opening. The process of both molts is undergone by the 
females while the hypostome is firmly inserted in the skin of the host. 
The shed skin splits open along either side and drops off in two scale- 
like j)ieces. A portion of skin from the capitulum is also slied at the 



LIFE HISTORY OF CATTLE TICK. 29 

same time. In the case of males, while durini^ and after the first molt 
the tick remains fixed to the host, after the second molt it detaches 
itself and travels in search of a female. As a rule the males molt two 
or three days ahead of females and frequently two are found attached 
to the skin of the host directly beneath nymphal females, awaiting 
the molting of the latter. 

When the adult stage is reached the development is very rapid. 
In from as few as 4 to 14 days the females become fidly engorged 
and fall to the ground to deposit their eggs. They die when the 
operation is completed. 

Although female ticks are somewhat more easily removed at the 
time of molting than at other times, repeated careful observations 
show that they do not actually detach themselves at either molt. 
However, Mi*. B. H. Ransom has shown that, when detached arti- 
ficially just before or after the second molt, females will reattach if 
placed upon another animal." Likewise he found that ticks removed 
just after the first molt would reattach after 24 hours. Specimens 
detached just before the second molt transformed and lived without 
host for two weeks. Several experiments have led us to the con- 
clusion that only in the rarest accidental cases can reattachment 
normally take place. In many cases we have attempted to cause 
tletachetl ticks to reattach, but in only two cases (hd we succeed. 
The following are some of the particulars: 

February 23, 1906, eight ticks, ranging in size from nymphs to one- 
half engorged adults, were placed on the shoulder of a steer. The 
next day all but one of these ticks had disappeared. One, however, 
wliich was about one-half engorged, had fastened to the skin about 
6 inches from the point where it was i:>laced. TMs tick was found 
detached on March 1 and trying to crawl out from the hairs which 
had been glued together to hold it in place. At tliis time it was not 
fully engorged. It began depositing eggs on March 18, and con- 
tinued oviposition until April 26, reaching a total of 523 eggs. An 
attempt was made to cause these eggs to hatch, but without success. 
It is not likely, however, that the failure to hatch was due to the 
experience of the tick. It is probable that the state of engorgement 
may have had sometliing to do with the matter, and, moreover, at 
the time of the year when the eggs were under observation it is 
exceedingly difficult to cause them to hatch. After the experiment 
that has been mentioned repeated attempts were made to obtain 
other cases of reattachment. Ticks at various stages were placed on 

« Lahille (Contr. 1' Etude Ixodides Ai-gentine, p. 112) had previously detailed a num- 
ber of experiments in the reattachment of Margaropus {Boophilus) microplus. It was 
found that immediately after molting the ticks would more or less frequently reattach 
after being removed artificially. Nothing, however, was found to indicate that the 
parasites naturally detach and reattach on the host. 



80 



NOETH AMERICAN FEVER TICK AND OTHER SPECIES. 



the steer and confined to limited area by means of vaccination 
shields. In no case under this manipulation did reattachment take 
place. Recently, however, a tick removed just before the second 
molt became adult in a pill })ox and reattached after 25 hours. 

ADULT STAGE. 

Adult females are the ticks that are generally seen, and their 
appearance is familiar to most persons. The males (which do not 
become engorged) are generally overlooked, although they may be 
easily found attached to the skin of the host direct h- beneath the 
females. This gives rise to a rather prevalent popular idea that 
females carry young with them. 

The following descriptions are taken from the work of Salmon and 
Stiles:" 

Afale.—liody ..val. naiTowed „n Innit, bn.adost (J. 3 inin.j at s^iigmal plane, 2.15 to 
2M mm. long. Scutum reddish ')rown, covering entire dorsal surface, prolonged in 
front by two pairs of i^rojections— one pair of more prominent dorso-lateral projections, 
dorsal of anterior projection of coxas I, and one pair somewhat less prominent and 
more median, ventro-median of first pair and nearer the neck. Two cervical furrows 
shallow, extending more or less distinctly 1o the posterior border; may be somewhat 
interrupted in the middle; a median furrow present in posterior half, may be very 
indistinct; posterior margin of body divided into festoons, which mav be only slightly 
marked. Relatively large circular pores, witli extruding .short bristly hairs, scattered 
over entire surface. Eyes small and pale, often problematic, at I intercoxal space 
Ventral surface lighter tlian dorsal, all portions provided with short stout hairs; geni- 
tal pore, broad, transverse, between coxte II; anus slightly posterior of stigmal pUine- 
(wo pan-s of anal plates (clypei): one pair elongate, rectanguhu- lo triangular, close to 
anus, in some cases extending cephalad to middle of coxa- IV. and caudad to near or 
beyond posterior margin, tlie anus being about at the middle of the length, in other 
(;ases extending from height of middle of stigmal area to beyond posterior margin of 
body; the median border longer than lateral border, the former prolonged into a point 
posteriorly, the postero-lateral margin may be nearly straight, or somewhat curved, or 
irregular in outline, thus presenting broad tooth-like projections; lateral and contigu- 
ous to each of these shields is found another shield somewhat similar in form, but 
smaller in size. Median caudal appendage ab.sent. Capitulum 450 to 500// long, ' base 
similar to that of the female, but a little straighter, longer, more salient in front of 
dorsal shield, into which it penetrates by a sort of rectangular neck, lateral projec- 
tions not very prominent. Mandil>les mOn long, digit about 90//; internal apophysis 
with straight base and broad iMfuh' point; external apophysis bidentate, the terminal 
subventral tooth may be very small while the proximal tooth is strong and large, or 
both may be large. Hypostome similar to that of female, four distinct rows of tee'th on 
each half. Palpi about 190// long, similar to those of female. Legs strong; coxae large, 
those on each side contiguous, as broad as long; coxaj I triangular, apex may be pro- 
longed anteriorly beyond the corresponding anterior point of dorsal shield, reaching 
anterior angle of base of capitulum, or may be very short, base posterior and more or 
less distinctly bidentate, the teeth short, often slightly pronounced, or quite promi- 
nent, the lateral tooth in .some cases prolonged into a well-marked spine. Tarsi like 
those of female. 



" " The cattle ticks (Ixodoidea) of the United States: - 17th Ann. Rept. Bur. Animal 
Industry, U. S. Dept. Agric, pp. 420-424, 1901. 



LIFE HISTORY OF CATTLE TICK. - 31 

Female. — Body elliptical, as broad in front as in back, usually somewhat con- 
stricted in middle, near IV pair of legs; may attain 13 mm. long, 7.5 mm. broad. Color 
exceedingly variable; live specimens vary from a tawny yellow (younger forms) to an 
olive green (very old specimens), alcohol specimens from yellow to red or black; the 
excretory system often shows tlu-ough the cuticle as tortuous whitish canals. Dorsal 
shield (scutum) very small, visible as a dark brownish spot in a depression at anterior 
end of median line; usually about 1.1 mm. long by 0.8 to 0.9 mm. broad, decidedly 
emarginate anteriorly to receive capitulum; lateral borders nearly straight and par- 
allel- in anterior portion, from antero-lateral points to eyes, then convergent from 
eyes caudad, forming a more or less bluntly rounded posterior angle in median line; 
cervical grooves 'divide the anterior half of scutum into three more or less equal lon- 
gitudinal fields, and diverge posteriorly; surface of scutum provided with short bris- 
tles which are more numerous near anterior border and near the eyes than elsewhere. 
Eyes rather small near anterior third of margin of scutum. On nearly the entire 
length of dorsal surface of body are two antero-posterior grooves, interrupted or nearly 
effaced near plane of IV pair of legs, and ending a short distance from the shield and 
from posterior margins of body; also, an unpaired median groove in posterior half of 
body; all three vary with the muscular contractions and may more or less com- 
pletely disappear when body is replete. Ventral surface shows four pairs of mon; 
or less distinct marginal constrictions corresponding to the four pairs of legs, the IV 
constriction being most marked, antero-median region also depressed at insertion of 
capitulum; vulva small, median, at plane of coxae I; sexual grooves corresponding 
to the paired dorsal grooves; but showing some variation in different specimens; 
median groove extending from anus to posterior margin. Anus about on border of 
second and last thirds of body. Stigmata short oval; stigmal pore slightly crescentic, 
convexity lateral; stigmal field with numerous larger and smaller wart-like structures, 
forming a zone near the margin. Cuticle of entire body finely wrinkled, liearing short 
hairs. Capitulum very short, about 800// from posterior dorsal margin to anterior 
end of hypostome; base of capitulum hexagonal, enlarged on its dorsal surface: in- 
serted in emargination of scutum; lateral projections not very prominent. Mandibles 
860/^ long, digit 120//. Internal apophysis conical (Neumann), bidentate (Fuller), 
with its base near the terminal extremity; external apophysis with tlu-ee successive 
teeth, one terminal, subventral, small; the second stronger; third large. Hypostome 
rather spatulate, broad, a little longer than the palpi, provided on each half with 
four rows of nine to ten nearly regular denticles, which do not extend to the base. 
Palpi very short (310/0, subconical, articles at least as broad as long; first article par- 
tially hidden under the antero-dorsal border of the base of the capitulum; second 
article pedunculate, dilated in a salient crest in its middle portion, thus forming a 
prominence inward (toward median line) and outward, and provided with strong 
hairs, especially on the inner prominence; third article smaller, subtriangular on its 
dorsal surface, where it forms a projection in and out; fourth article small, cylindrico- 
conical, infero-terminal. Legs rather thin, short (pair I, 2 mm.; pair II, 2.5 mm), yel- 
lowish brown, first articles darker than the others. Coxae: pair I subtriangular, pos- 
terior border bidentate or biundulate, the division in many cases indistinct. Tarsi 
I unicalcarate, II to IV bicalcarate. Pulvillum about half as long as claws. Stiff 
l)ristle-like hairs on all articles. 

EFFECT OF CONTINUOUS COLD AND HEAT ON ENGORGED 
FEMALES. 

Twenty-five ticks were used in the experiments with cold, which 
were conducted durmg the month of August, 1906. A mean tem- 
perature of about 48° F. was maintained, with extremes ranging 



82 



NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 



from 34 to 53 _. M ith exposure up to 300 hours practically all ticks 
recovered and m most cases deposited viable eggs. In cas;s of more 
than 300 hours exposure practically all ticks survived, but none 
deposit(Hl via})le eggs, although in many instances oviposition took 

JV ?noo^'' ''^ ^-^P^""^^"t« with heat a mean temperature of from 
J8 to 102 was mamtamed. Up to 103 hours of exposure to this 
temperature practically all ticks deposited eggs that were viable 
With exposure at the same mean temperature of from 144 to 21S 
hours duration, eggs were deposited, but were found not to be viable 
1 hey were dry and shriveled when deposited 

Some of the females survived heating for the h.ngest period, namely, 
218 5 hours \Vith an exposure of 103.5 hours or more, however, a 
least one-half succumbed. 

EFFECT OF DIRECT SUNLIGHT ON ADULTS. 

Eleven unengorgcd fcn.ales placed i„ a box exposed to the direct 
^- of the ju,n m SeptemW died in three days. Seven unmatured 
females m dirc^c sunhght from morning until noon seemed dead at 
noon. They did not survive until the next dav, although they were 
removed from the direct sunlight at 2 o'clock." Similar experiments 
showed that death resulted in the case of engorged females after a few 
hours exposure to the sun. In experiments with eggs, tubes were 
sul,jected to direct sunlight for one day. Wlu>n n.oistened while 
kept^ ui these tubc^, hatching seems to take place normally, and 
hatching foll,>w(Ml m similar experiments in ^vh\v}x the e-o-s were 
kept dry. '^'^ 

EFFECT OF SUBMERGENCE IN WATER ON ENGORGED ADULT 



TICKS. 



Adult ticks have remarkable resistance to the elFect of submergence 
as has been pointed out to be the case with eggs and seed ticks. The 
iinmediate effect of submergence is to cause a cessation in the activ- 
ity of the ticks, while they become somewhat distended apparently 
from the absorption of water. In August and September, 1905 a 
considerable number of experiments were conducted in which" the 
adult ticks were submerged in water from the city mains at Dallas 
I ex. Judging by the experiments with seed ticks and eggs mentioned 
elsewhere it is not likely that water impregnated with foreign matter 
would have changed the results. During the months mentioned a 
period of submergence of 24 hours did not result in the death of any 
appreciable number of ticks used in repeated experiments After 
one or two hours the specimens recovered from the immediate effect 
of submergence and proceeded to deposit eggs which were found to be 



LIFE HISTORY OF CATTLE TICK. 38 

viable. With between 24 and 48 hours' submergence the number of 
ticks that recovered diminished rapidly. Occasionally specimens 
recovered after a period of submergence of over 48 hours. For 
instance, the specimen collected on July 12 and submerged for 50 
hours survived and deposited viable eggs. Later in the season (that 
is, in October) somewhat difl'erent results were obtained in experi- 
ments in the submergence of adults. In this month many ticks 
recovered after from 50 to 90 hours of submergence. In fact, fully 50 
per cent of the ticks submerged between these extremes regained 
their fidl activity. In one experiment two out of five ticks sub- 
merged in October for 91 j hours recovered and deposited viable eggs. 
Nevertheless a number of ticks submerged for 115| hours did not 
recover. 

The results that have been mentioned above indicate that where 
engorged ticks fall from cattle that are standing in permanent pools 
of water none will survive to deposit eggs. At the same time the 
results show that temporary flooding of from 24 to 100 hours' duration 
would not in all cases prevent ticks from depositing eggs. It must 
be noted in this connection, however, that the vitality of eggs depos- 
ited by ticks just prior to temporary flooding would not be interfered 
with by the water, although, of course, they might be washed away. 

DROPPING FROM HOST. 

It is a more or less prevalent popular idea that ticks have some 
sense which enables them to drop from a host in places favorable 
for oviposition. A few observations have shown, as was supposed in 
the beginning would be the case, that there is probably no such 
power of perception present in the cattle tick. The popular impres- 
sion probably had its origin in the fact that bunches of seed ticks are 
found most numerously in the places where the cattle collect for the 
benefit of shade or water. This phenomenon obviously is due merely 
to the collecting and standing of the cattle. 

Many observations seem to show clearly that there is no preference 
as to the time of dropping. In our experiments many ticks have 
been known to drop from host during the night and many others to 
drop during the day. 

LOCOMOTION. 

Some attention has been paid to obtaining data on the distance 
engorged ticks may travel after dropping from the host, since tliis 
has a practical bearing on double fencing in eradication work. In 
some experiments specimens w^ere placed on the floor in the laboratory 
and allowed to move at will. The total movement varied from 24 
inches to 123 inches, the latter distance being covered in 52 minutes. 
5795— No. 72—07 'd 



34 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

The effect of antiheliotropism was always in evidence in these exper- 
iments. The hght was admitted from different quarters at (hfferent 
times and the ticks ahvays changed their course so as to travel awa}' 
from it. These results are very similar to those published by Lahille 
from experiments with Afargaropus (Booj^hilus) microplus in Argen- 
tina. The distance engorged females will travel depends upon how 
soon they can reach shelter. If the}^ obtain protection under debris 
of any kind the}'^ seem to be disinclined to go farther. The greatest 
distance traveled on the ground out of doors was only 24 inches. 

It is popularly supposed that engorged female ticks often burrow 
into the ground. In a nund^er of experiments with light barnyard 
trash in tubes it was found that females will work their way down 
from one-half to IJ inches. In such cases the masses of eggs assume 
various shapes on account of the surrounding debris. (See PI. I, 
fig. ].) It was found that seed ticks from eggs so deposited had no 
difficulty in reacliing tlie surface. 

HOST RELATIONS OF THE CATTLE TICK. 

Margaropiis nnnvlatus was descril)e(l in 1S21) by Say "'from speci- 
mens taken on a Virginia deer in Florida." Since that time we 
have been unable to find records of the occurrence of the species on 
deer. However, a nundjer of instances have come to light recently 
in which undoubted specimens of Margaropus annulatus have been 
found on deer. The first of the.se cases was the result of an examina- 
tion made by Mr. R. C. Howell on a herd of tame deer in a park at 
Mount Pleasant, Tex., in October, 1905. Since that time Mr. J. I). 
Mitchell has found specimens on a deer at Oakville, Tex. Mr. T. R. 
Coker lias sent specimens from the same locality and host. In both 
of these instances the ticks were found on wild deer that had just 
been shot: In February of the prescmt year Mr. F. C. Pratt collected 
a few sj)ecimens of Margaropus annulatus from a dry deer liide at 
Kerrville, Tex., and in December he examined a fresh liide on which 
considerable numbers were found. 

The matter of the possible develoj^ment of Margaropus annulatus 
on various animals, among them guinea pigs, rabbits, dogs, and cats, 
has been studied by various investigators. Dr. J. W. Connaway's 
experiments in ^Missoin'i in 1897 showed that the ticks would not 
attach to rabbits or guinea pigs. They did attach to dogs in consid- 
erable numbers, but only one of them ever matured. Recently 
Air. B. II. Ransom has repeated the experiments with rabbits, dogs, 
and cats. On rabbits and dogs the ticks attached, but remained so 
only a short time. On a cat a female tick remained attached from 
July 25 to August 30 and molted on the host. Nevertheless, it did 
not reach engorgement. 



HOST RELATIONS OK CATTLE TICK, 85 

In our experiments with ticks we have kept several dogs primarily 
for experiments with species other than Margaroims annulatus. At 
the same time we have repeatedly sprinkled thousands of seed ticks 
of Margaroims annulatus on these dogs, but in no case have we noticed 
that attachment took place. This, with the work of Comiaway, 
Schroeder, and Ransom, seems to indicate that dogs can play but a 
very unimportant part as hosts for the cattle tick. Ransom men- 
tions the fact that the collection of the Bureau of Animal Industry 
contains specimens of Margaropus annulatus collected from a dog, 
from which host Francis (1894) seems to have been the first to report it. 

One of the writers has had one case of an attachment of Margaropus 
annulatus to his person. This was a male specimen that attached 
between the fingers of the hand. It was removed after about half an 
hour. Mr. Ransom mentions a similar case in which, however, the 
specimen was a female and remained attached to his hand for twenty- 
four hours before it was removed. Attachment to human beings must 
be very rare. The junior author had worked with thousands of 
specimens of ticks before and after the single case of attachment that 
has been mentioned. 

In addition to the abnormal and unusual host relations mentioned 
above, there are not infrequent instances in which Margaropus annu- 
latus has been found to occur on horses, mules, and asses. 

All this work shows clearly the remarkable host restriction of the 
cattle tick that is most important from the viewpoint of its attempted 
eradication. 

The early records of Packard, showing the occurrence of the cattle 
tick on a porcupine and a similar record of its occurrence on the 
rabbit in Idaho, must have been due to a misidentification of the 
species. 

The strict instinct for the proper host in the cattle tick is shown in 
the extreme infrequency of attachments of ticks to each other. 
Thousands of ticks have been sent to the laboratory alive inclosed 
together in tin or wooden boxes. In only one case was it found that 
a tick had inserted its rostrum in another. This happened in a lot 
collected in southwest Texas by Mr. J. D. Mitchell on April 19. A 
female had inserted its beak firmly on the lateral dorsal portion 
behind the posterior coxse. The two specimens were placed in 
alcohol and still remained connected." 

RELATION BETWEEN RATION AND TICK INFESTATION. 

For several months after experiments were started at Dallas in 
placing seed ticks on the steer used for experimental purposes it was 
found that a surprisingly small proportion ever became adult. In 

"Since the above was written important discoveries regarding the occurrence of 
Margaropus annulatus on sheep have been made by the Bureau of Entomology. See 
Circular 91, Bureau of Entomology, U. S. Departmentof Agriculture, issued July 3,1907. 



3(3 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

man}^ cases from 5,000 to 10,000 seed ticks were placed on the animal, 
but only from 1 to 20 adults ever developed. Prof. H. A. Morgan 
made the suggestion, based upon the observations made by him in 
Louisiana, that the ration the steer w^as receiving was responsible for 
this remarkably small proportion developing. At his suggestion we 
changed the ration. The steer had been receiving daily 4.76 pounds 
of corn chops and 5 pounds of prairie hay. The inspection tag on the 
chops guaranteed not less than 9 per cent protein crude and not less 
than 4 per cent fat crude. At Professor Morgan's suggestion the 
corn chops were eliminated. Immediately a much larger percentage 
of seed ticks developed to adults on the animal, although his general 
condition did not seem to have been changed materially. While 
before a dozen adults from many thousand seed ticks was the maxi- 
mum, after the change in the ration hundreds developed fr<^m no 
larger numbers of seed ticks applied. 

ENEMIES OF TICKS. 

At one time it was supposed that sowbugs may sometimes be 
im})ortant factors in the destruction of tick eggs. A number of 
observations have shown that the greatly preferred food of these 
isopods is vegetation either live or decayed. In laboratory experi- 
ments Armadillidium vulgare was found to feed on dead ticks and 
also to devour the eggs whenever no other food was provided. Thirty- 
eight sowbugs, furnished with 897 tick eggs, consumed 366 at the rate 
of 3 eggs per day each. In another case two sowbugs devoured 159 
tick eggs at the rate of 15 each per day. These results hardly seem to 
substantiate the impression that sowbugs may be of considerable 
economic importance. It should be emphasized that the experiments 
referred to were conducted in the laboratory, and the sowbugs were 
deprived of other food. Under natural conditions the results might 
have been quite different. 

The little "fire-ant" (Solenopsis geminata Fab.), which has recently 
been found to be acquiring a special taste for the boll weevil, undoubt- 
edly destroys many engorged ticks that have dropped to the ground. 
Experiments performed by placing ticks in the immediate vicinity of 
nests of this ant show that under such circumstances they must 
invariabl}' be killed. The nests of this ant are found throughout the 
pastures in the South, and the total of the work done by them must 
be considerable. 

A number of dipterous larva? have been found feeding upon tick 
eggs and an undetermined species of Phorida^ has been bred. 

At one time we were inclined to believe that a chalcidoid parasite 
of the cattle tick had been reared. Early in 1906 such a parasite 
was found in a pill box with the remains of an engorged tick placed 
there the fall before. Upon sending the specimen to Dr. L. O. How- 



ENEMIES OF TICKS. 37 

ard it was learned that it belongs to a new species and creniis. Chalci- 
doids of the subfamily to which it undoubtedly belongs are known to 
be parasitic on various dipterous larvse. Upon reexamination of the 
remains of the tick a portion of a dipterous cocoon was found. Con- 
sequently the hymenopteron was probably not a parasite of the tick, 
although the interest remains, since the dipteron was probably para- 
sitic on the tick. 

A number of observations have been made showing that domestic 
fowls frequently learn to remove ticks from cattle in barn lots. Mr. 
F. C. Pratt observed a case in which the fowls regularly visited hides 
hanging up to dry for the purpose of picking up the ticks which 
dropped from them. Mr. J. D. Mitchell has witnessed '" jackdaws " 
(Qaiscalus major macrourus) picking ticks from cattle near Victoria, 
Tex., and the farmer informed him that he believed these birds kept 
the cattle practically free of engorged or nearly engorged specimens. 
Mr. S. E. McClendon, of the Louisiana Experiment Station, informs 
us that he has repeatedly seen kingbirds ( Tyrannus tyrannus) engaged 
in the same work. 

In connection with the enemies of ticks it may be stated that it 
seems likely that mice are of some importance. In the laboratory 
it was found that the best bait for mouse traps that could be used was 
engorged cattle ticks. It seems likely that in the pastures field mice 
may frequently devour ticks. Button and Todd (Human Tick fever, 
p. 17) write as follows: "Ticks are not without natural enemies. 
Rats eat adults with avidity, and ants carry off young ones and 
eggs. We have lost ticks in both ways. One occasion over two 
hundred young ticks were carried off in a single night by small 
ants." These remarks apply to OrnitJiodoros mouhata in West Africa. 

THE PRACTICAL APPLICATION OF THE INFORMATION RECORDED 
IN THIS BULLETIN. 

In the preceding pages at difl'erent places the special practical 
importance of the data discussed has been mentioned. As a matter 
of fact the work upon which this bulletin is .based has been planned 
to accumulate additional information for use in the practical work of 
tick eradication. Some methods of control are satisfactory in certain 
districts, but much less so in others. Plans that would be feasible 
along the northern border, for instance, where the tick is on a rather 
delicate equilibrium and is never found on the cattle for months during 
the winter, would not be applicable to the moist regions along the 
Gulf where the cattle are infested throughout the year. Of the 
various methods of eradication undoubtedly those of the widest 
utility are the ones which prevent the development of the tic4cs by 
breaking up the relation between them and the cattle. Of these, the 
more important are the feed-lot or soiling system for relieving the 



38 



NORTH AMERICAN B^EVER TICK AND OTHER SPECIES. 



cattle of ticks and the pasture-rotation system (to be used in con- 
junction with the former) for freeing the pastures. 

In the feed-lot or soiling system the basis is the time occupied for 
development on the host in connection with the time from the drop- 
ping of engorged females to the hatching of seed ticks from eggs 
deposited by them. In a tick-free inclosure, as a feed lot, cattle 
may be left to drop their ticks until it is time for the eggs from the 
first-dropped individuals to hatch. Information as to exactly when 
it will be necessary to move the cattle to avoid reinfestation is given 
in Tables I and II, which show the numbers of days at different seasons 
before ticks begin to oviposit after dropping and the time before the 
first eggs deposited will hatch. In July, 1906, for instance, this was 
from 25 to 27 days. The time required for all ticks to drop from 
cattle, which indicates how long they must remain in one or more, 
tick-free areas, is shown in Table VII. The period is from 31 to 59 
days. 

The data necessary for an intelligent plan of freeing pastures, by 
removing the cattle until the death of the ticks, are given in Tables 
V and VI, showing the period from dropping to the death of all the 
resulting seed ticks. The former table shows tiiat if cattle were 
removed from a pasture on June 20, 110 days later it would be per- 
fectly safe to consider it tick-free. 

The data referred to above, together ^^^th the results of other 
experiments, have been arranged to cover maximum preoviposition 
and maximum oviposition periods, for convenience, in Table VIII. 

Table VIII.— /'(7-/o(/.s^ in the life hif<tury of the rattir tick upon which means of control 

are based. 





Period from dropping 
to oviposition. 


Minimum 

incnliMlion 

period. 


Minimum 
l)eriod from 
dropping of 
ticks to 
hatching 
of eggs. 


Seed ticks 
all dead. 


Maximum 
period from 
dropping of 
adult to 
death of 
seed ticks. 


When licks dropped. 


Minimum. Maximum. 


1905. 

Sept. 2 

Do 

Dec. 23 

Dec. 24 

Doc. 25 

19011. 

Mar. 20 

Apr. 19 

Apr 21 


Dni/s. Da II It. 

3 " 5 

3 5 
21 41 
21 41 
21 41 

9 10 
5 6 

5 

6 ' 10 
6 10 

4 6 
4 6 
3 5 
3 , 5 
3 , 5 

l\ . 5 

3 4 

V\ \ 

3 i 4 

I 


54 
47 
47 

39 
33 
29 
27 
28 
27 
27 
28 
23 
23 
23 

22 
23 

24 


^''^%6 
59 
75 

()8 
08 

i 

34 


June 22 
May 31 
Aug. 28 
Aug. 15 
Aug. 22 

Aug. 28 
Sept. 20 
Oct. 5 


Days. 

293 
271 
248 
234 
240 

161 
154 
167 


Mavi 


33 '....do... 


157 


May 4 

May 19 

May 21 . 


34 
31 
31 
31 
26 
26 
26 
24 
25 

. 1 

27 


Aug. 9 
Sept. 20 
Sept. 22 
Aug. 15 
Aug. 26 
Aug. 6 
Oct. 6 
Sept. 26 
...do... 
Oct. 20 
Feb. 18 
Jan. 23 


97 
130 
124 


Junes 

June 4 


73 
a83 


June 6 


a 61 


Do 


122 
106 


June 13 

July 1 

July 13 

Do 


105 
olll 
«220 

194 



; These ticks were kept in open-bottom test tubes exposed to sun at all times. 



APPLICATION OF INFORMATION. 39 

In the above experiments (Table VIII) the og,^s and seed ticks, 
except those marked ("), were kept in cyhnch-ical tiilx's of l-incli 
diameter inserted for about Ih inches into sand or- soil in receptacles 
on or below the surface. These were kept in a cage on the west side 
of the laboratory so that they were protected from the sun until 11 
a. m. As soon as the eggs commenced to hatch, there was jilaced in 
the tube a stopper of absorbent cotton which remained unremoved 
until the death of the ticks. In cases marked (") the conditions were 
the same, except that the tubes were exposed to the sun at all times. 

The conditions furnished included a humid atmosphere and a min- 
imum temperature as compared witli the normal. The enforced 
bunching and protection from wind and rain furnished additional 
favorable conditions. It would seem that the periods are maxima 
and hardly to be met with in the work of extermination. One unfa- 
vorable factor met with, which could not be prevented, was the 
growth of algse on the inside of some of the tubes. This accounts for 
some of the variation in the longevity. 

In addition to the main practical data given the following are of 
importance: 

All exj^eriments and observations show the close host restriction 
of the cattle tick. Though the occurrence of the cattle tick on deer, 
horses, mules, asses, and some other animals is comparatively rare, 
it must be taken into consideration in the i:)ractical work of tick eradi- 
cation. 

Eggs are but little affected by water. When submerged, they 
hatch in about the normal time. Seed ticks are also resistant to water. 
In one case seed ticks survived submergence of 157 days. Adults 
are less resistant, but in the summer they frequently survived sub- 
mergence of 48 hours. Later in the season the resistance seemed to 
be greater, some females surviving after more than 90 hours. It is 
evident that water courses must play a very important part in tick 
dissemination. 

The adult cattle tick has only the most limited means of locomotion. 
After it drops from the host it crawls but a few feet at the most. 
On the ground in our experiments the greatest distance traversed 
was only 24 inches. 

On the whole, no very imiDortant enemies of ticks have been found. 
Domestic fowls frequently devour considerable numbers of them, and 
some wild birds also render valuable assistance in picking them off 
animals or from the ground. 



NOTES 0\ \ ARIOUS SPIXIES ()V TICKS FOUND IN THl: 
UNITFT)STAT1:S. 



The role that Margaropus aintulatus Say was found ])y Smith aiul 
Kilborne to play in the transmission of splenetic or Texas fever in 
cattle impressed upon investigators the importance of ticks as carriers 
of disease. Since that time the study of these creatures has progressed 
rapidly. Smith and Kilborne, Lounsbury, Theiler, Marchaux, Salim- 
beni, Dutton and Todd, Motas, Kossel, Ricketts, and King are among 
those who have demonstrated that ticks are the agents through which 
various diseases of man and other animals are transmitted. What 
is greatly needed in this country at the present time is a convenient 
means of identifying the various species. This the writers have 
attempted, in a measure, to supply in the following pages, in which 
will also be found notes on the life history and habits of such species 
as they have encountered. 

CLASSIFICATION AND HABITS OF TICKS. 

The following key will (Miablc one to determine the genera of the 
various ticks found in this country: 

KEY TO FAMILIES, SUBFAMILIES. AND NORTH AMERICAN 
GENERA OF TICKS, i IXODOIDEA i. ' 

Scudiin al)sent Family Ai;<i asid.k. 

Scutum present .Family Ixodid.k. 

Family ARGA.SID.E. 

(.'apitulum at least its Icntrth from the anterior margin Genus Arga.s ( p. 42). 

C'apitulum under a l)eak-likc projection, close to anterior margin. 

Genus Ornilhtxloros (p. 45). 
Family IXODIDJv 

1. Palpi shorl. nol m- cuily .-^ligluly longer than liroad: cajjitulum short. 

Subfamily Rhipiceph.vlin.e, 2 
Palpi ])lainly longer llian hniad; capituluni longer Subfamily I.xodin.e, 5 

Sul)family Rhipickphai.in.k. 

2. Dorsal surface of capituluni hexagonal, the sides projecting in angles; male with 

anal plates 3 

<' This table is based ui)on those of Salmon and Stiles (1901) and Banks (1904). The 
genus Ceratixodes is not included (see p. 54). 

40 



CLASSIFICATION AND HABITS OF TK^KS. 



41 



Dorsal surface of capituluin rectangular, sides straighl; male without anal plates. A 
?>. Second and third palpal segments extend laterally into sharp points; stigmata 

nearly circular Genus Mnrgnrnpvs (p. 40'). 

Second and third palpal .segments even: stigmata comma-shaped. 

Genus Rhipiccp/irilus (p. -17). 
4. Eyes present; external hordor of ])alpi straight; cox:e I Indcntatc. 

Genus Ddiuaccnior (]>. 49). 
Eyes absent; external l)or(U'r oi palpi uneven; coxje I not l)idciilalc. 

Genus //.riiKiphi/sdlis (p. 52 K 
Subfamily Ixodin k. 

o. Anal groove surrounds anus anteriorly and opens jjostcriorly; eyes absent; stigmal 

l)late nearly circular Genus Ixodes (p. 54). 

.\nal groove surrounds anus posteriorly and opens anteriorly; eyes present; stigmal 
plate reniform Genus Awblynmma (p. 58 ). 

Lahille has recentl}'" published an inoenioiis gra])hic tal)le for the 
separation of the famihes and genera of Ixo(k)idea. Tt is reprodticod 



Familias 

Argasidae (^ 



SUB-ORDEN: ArPAGOSTOMA 

"c^'^.'^^ - Argas 

(X nrnithodoro5 - 

<^ HagmaphusaliS" 

^ -JW Aponomma " 

^ J*% _ Neumanmeiia 



lxodid«( 



^ I ^,^ Ambluomma 



Art^opliL^^ . ., 



O 
Anopli po-^^^^^i 

Ommataj C^M^ [)pf mdcenlof ' 

|j)^...„_^- HLjalomma 

mh Rhipicephalus " 

'^^- 1 ^' \^ t^ @----Boophilus" 

Penssoplij O I ^ -•■■ ■ "r^" . , 

.g^% Anoi^ata W. L schalocephalus 

Fig. 2. liraphictable for the separation of tlic familu>s and genera of ticks. (From Lahille.) The 
underscored genera ato represented in the United .States. 

here as figure 2. The genera underscored are known to be repre- 
sented in the United States. The suborder Arpagostonia of Lahille 
is the same as the superfamily Ixodoidea of Banks, used by us. 

Family ABGASID-ffi. 

The family Argasida^ is represented by two genera, Argas and 
Ornithodoros. They are readily distinguished by the characters 
given in the table. The species are parasitic on mammals and birds. 



olxodides Argent., 1905, p. 21. 



42 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

Genus ARGAS. 

The species belonging to the genus Argas are nocturnal parasites 
of chickens, pigeons, and other birds, and occasionally attack mam- 
mals, man included. Two species are represented in this countr}^ — ^4. 
miniatus Koch and A. sanchezi Duges. The former has been found 
by Marchaux and Salimbeni to transmit spirillosis of fowls in -Brazil. 
There is some evidence of the occurrence of this disease in the 
United States. 

ADOBE TICK. 

i Ar<ia.'i sanchezi Duges.) 

This species was described in 1891 b}' Duges from larva^ collected 
at Guanajuato, ^lexico, on a dove (Zenaidura macroura) . Tn struc- 
ture it is very similar to A. miniatus. Neumann's description is 
based upon a number of specimens collected at Mariposa, Cal., ui)on 
a (juail, and at Santa Agueda, Lower California, upon a wild turtle 
dove. Mr. Nathan Banks has specimens from Deming, N. Mex., and 
from Arizona, and it is (piite j)r<)bable that it will be found in Texas, 
and possibly farthei- north, ^^r. Banks states that in New ^Mexico it 
is found in houses and is there known by tli(> common name used 
above. 

KOWl. TICK. 

{Art/da iiiiuidliis Kocli.)" 

This species was described in 1S44 by Koch from Demerara. The 
name Argas americanus, applied by Packard in 1873 and used largely 
by writers in this country, is a sj^nonym of A. miniatus. For excel- 
lent illustrations of this species see Salmon and Stiles, Ixodoidea of 
the United States, Plate LXXII and text figures. 

The species is well distributed over the world. The records include 
Persia, ^Vlgeria, Russia, India, Australia, South Africa, Central and 
South America, and, in the United vStates, Florida, Texas, New 
Mexico, Arizona, and California. In southwestern Texas it is found 
in large nund)ers in and alxmt chicken houses and out-door roosts, 
hiding away in crevices by day and coming out at night to engorge 
on the fowls. This species seems occasionally to attack mammals, 
as Mr. J. D. Mitchell has taken it once in Texas from the rabbit. 

As pointed out b}" Lounsbury, the sexes are so much alike that, 
except by the size, the onl}^ safe way of separating them is through 
the form of the genital orifice. To quote from him, "This orifice 
[genital] is situated just behind the mouth parts on the under side of 
the front of the body; that of the male is relatively inconspicuous 



a The name of this tick is in great confusion. The one used by the writers is that 
recently adopted by Banks. 



CLASSIFICATIOlSr AND HABIT8 OF TICKS. 



43 



and is surrounded by an oval ring, and that of the female appears 
as a transverse slit in the leathery surface. Both sexes are of the 
same dimensions when they become adult. The male, however, does 
not perceptibly increase in length and width by feeding, whilst the 
female does, and hence amongst specimens collected about a fowl 
house the females are generally larger than the males." 

Its life history has been worked out by Lounsbury at Cape Town. 
As brought out by him, the life cycle and habits are strikingly dis- 
similar in some important respects from those of the true ticks, the 
general habits being found much more like those of the bedbug 
(Cimex lectularius L.) As might be expected, we have found the 
stages in the life cycle to vary somewhat from those he found. Unlike 
the ticks of the family Ixodidse, this species feeds for but a few hours 
at a time, and then always at night, excepting in the larval stage, 
when we have found it to remain attached for five to eight days 
before dropping. There is also an extra nymphal molt. Unlike the 
ixodid ticks again, these do not die following engorgement, but live 
on, ovipositing repeatedly. Within a week or ten days from one 
feeding in warm weather they again find a fowl and engorge. As 
many as five different feedings as adults have been recorded by Louns- 
bury, each followed by the deposition of eggs. 

In our experiments seed ticks were placed upon a fowl and obser- 
vations made to determine the period of larval attachment. In 
about three days from attachment they became rounded and black 
from the engorgement of blood. A few hours before dropping they 
commenced to flatten and assume the typical Argas shape. Attach- 
ment continues for from five to eight days. In September and Octo- 
ber fourteen days were found to pass after dropping before molting 
took place. The attachment for second and third engorgements 
Lounsbury found to last but a few hours, about two weeks to pass 
after the second, and a like period following the third engorgement 
before moltino-. 



Table IX. — Ovipnsitinn of Argas nriniatiis at Dallas, Tex. 



■ 


First engorgement 
recorded.- 


First oyiposit 


ion. 




When collected. 


From— To— 


Num- 
ber of 
days. 


ber of 
eggs. 


1906. 
May 12 




June 23 July 4 
May 17 | May 30 
May 16 May 23 
May 17 May 29 
May 18 June 3 
May 18 May 24 
May 16 May 22 
May 2 May 20 
Aug. 1 Aug. 10 


12 
14 
8 
13 

.19 
10 


113 
974 


Do 


do 


Do... 


do 


1.58 
169 


Do 


.do 


Do 


do 


194 


Do 


do 


50 


Do 






Mar. 24 


Apr. 17-18 




Do 


tin 


83 


■ "" 





44 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

Table IX. — Oviposition of Argas miniatus at Dallas, Tex. — Continued. 





Second 
engorge- 
ment. 


Second oviposition. 


Tliird 
engorge- 
ment. 


Total 


\Vhcncollcot<'(l. 


From— 




iNum- berof 
To— berof ^ggs- 
days. 


num- 
ber of 
eggs. 


190(;. 
Mav 12 








Do 

Do 

Do 


Aug. 8- 9 
Aug. 9-10 
.\ug. 10-11 
Aug. 15-16 
Aug. 17-18 
Aug. 23-24i) 
Aug. 7- 8 
Aug. 18-19 


Aug. 'U 
Aug. 15 
Aug. 16 
Aug. 21 



Aug. 24 , 11 180 
Aug. 21 7 199 
Aug. 25 10 193 
Sept. 4 15 1 148 



Oct. 22-23 
Sept. 3a 


454 
357 


Do 

Do 


Oct. 17-18 


342 
50 


Do 




32 


Mar. 24 

Do 


Aug. i7 
Aug. 24 


Aug. 24 8 ! 55 
Sept. 2 10 ' 154 


Sept. 28^29 
Oct. 16-17 


185 
237 



a Dead. b Last engorgement. 

Adult ticks wei^ihed before and after engorgement were found to 
increase in weight more than 300 per cent. 

In order to determine the incubation period, 35 daily lots of eggs, 
deposited between May 16 and September 1, were recorded. Of 
these, four lots commenced hatching in 14 days, 26 in 15 days, and 5 
in 16 days. In the incubator eggs deposited August 21 and subjected 
to a mean temperature of 99.8° hatched on August 29, the maximum 
temperature being 108°. From experiments carried out by placing 
eggs and seed ticks in an ice "box and exposing them continuously, 
these were found to be exceedingly resistant to cold. Eggs deposited 
August 27 were exposed from September 8 to October 1 to a mean 
temperature of 48.9°, a maximum of 67° and a minimum of 37°. 
These commenced hatching October 6. Two lots of larvae, one of 13, 
that hatched September 2, and a second of 30, that hatched Septem- 
ber 8, were exposed in the ice l)ox from September 8 to October 22 
to a mean temperature of 45.9°, the maximum being 67° and the 
minimum 36°. These were all alive when removed and were as active 
as ever October 25. 

At Dallas larva? kei)t submerged in water to a depth of about an 
inch lived for 1 1 da^s. 

The length of life of this tick and its capacity to exist in the absence 
of a host are surprising. At Dallas larva? ke])t confined in summer 
in pill boxes immediately after hatching lived about two months, 
some surviving somewhat longer. Larva? of Margaropus annulatus 
kept under similar conditions live for but two or three days at the 
most. In Australia Robertson found the nymphs to live in pill ])Oxes 
for about the same j^eriod as we have found the larvae to survive. 

The longevity of the adult, however, is most remarkable. Riley 
reports an adult specimen as remaining alive in a corked vial without 
food for five years." Robertson has found them to remain alive for 
two years and three months and Dr. Cooper Curtice informs us that 
he has kept them alive without food for more than two years. In 
our experiments adults collected in March, 1906, and kept in corked 



a Pr, 



Ent. Sue. Wash.. III. p. 121. 



CLASSIFICATION AND HABITS OF TICKS. 45 

vials are still alive, March 1 , 1907, althoii<i;h a number have succumbed. 
Not less surprising than the longevity of the adult is its resistance to 
insecticides. Lounsbury has kept adults confined for three months 
in a box nearly filled with flowers of sulphur with no apparent effect 
on them. He has also exposed them for two hours to hydrocyanic- 
acid gas at the strength of 1 ounce of potassium cyanid to 150 cubic 
feet of space and found that this scarcely served to decrease their 
activity. Further, many individuals survived for some days after 
treatment with parafFm and various oils. 

Genus ORNITHODOROS. 

Ornithodoros, the second genus belonging to the Argasida, is 
represented in the United States by two species, 0. )ue(/nini Dug., 
and 0. turicata Dug., both known to attack man. 

A species widely distributed through Central and South Africa, 
0. mouhata, was reported by Dr. C\ithbert Cliristy (1903), of the 
Liverpool School of Tropical Medicine, as the probable transmitter 
of tick-fever in man. In 1905 Dutton and Todd," not knowing of the 
work of the other investigators, demonstrated that "tick-fever" in the 
oriental province of Kongo Free State is a relapsing fever produced 
by a spirillum, probably Spirillum {S piracJisete) ohermeieri, and that 
this organism can be transmitted by the bite of the tick. 

The life cycle of members of this genus has yet to be followed. 
Lounsbury states that 0. savignyi begins to engorge at once when 
applied to a host, and that generally it is off again in an hour. After 
an engorgement, he states, it rests for many weeks or months and, gen- 
erally, at least, sloughs its skin if immature or lays eggs if a mature 
female before again seeking an animal. 

It is suspected by Mr. Nathan Banks that a species of the genus 
transmits a disease of cattle in California. 

SPINOSE EAR TICK. 

{Ornithodoros megnini Dugey.) 

This tick was first described in 1883 by Duges, from Guanajuato, 
Mexico, as a species of Argas. It has been reported in the United 
States from New Mexico, California, Kansas, and Nebraska, and is an 
important tick in Texas. The writers are mformed by a rancher in 
the western part of the State that considerable injury is due to the 
irritation produced by it in the ears of cattle and that its presence can 
often be told by the rough appearance of the hair. A prominent 
stockman in Dewitt County states that, in his opinion, it is second to 
the fever tick in importance. In addition to cattle it is found upon 
horses, asses, dogs, and sheep, and has been reported several times 
from man. Mr. J. D. Mitchell, of the Bureau of Entomology, reports 
two cases at Victoria, Tex., in which specimens were taken from 
human ears by a physician, following prolonged severe pain. Mr. 

"Memoir XVII of the Liverpool School of Tropical Medicine. 



46 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

Mitchell informs the writers that he has known this tick to climb 3 
feet from the ground and deposit its eggs in a crevice on the corner of 
a house and in other cases in cracks in the bark of trees at about the 
same height. From the action of specimens confined in nearly verti- 
cal vials he supposes that the species normally climbs some distance 
upward for the purpose of depositing the eggs. 

The nymph and adult are quite different in appearance, so much so 
that iN'eumann states that he would hesitate to consider the two forms 
as belonging to the same species had he not found a cast skin about a 
female. Marx figured the nymph as a new genus and species, Rliyn- 
choprium, spinosum . 

Stiles and Hassall have described and figured a pupa-like stage of 
this tick. This, however, does not seem to be a true resting stage, as 
specimens run very actively when removed and placed in boxes. The 
present writers, from live material in hand, are inclined to consider 
the so-called pupa-like stage as merely the engorged larva. 

The longevity of 0. megnini seems to be much like that of Argas. A 
specimen collected by the writers in July, 1905, lived to beyond Jan- 
uary 12, 1907, having been ke])t in a i)ill i)()x for a year and a half. 

TURICA'IA 1ICK. 

(Ornithodoros turinita Duges.) 

This tick was originally described from Mexico. It attacks man, 
the punctures causing large swellings that remain for several days 
and are followed by severe pain. It has been reported from South 
America on the llama; from the United States, in Florida, on a land 
turtle {Gopherus pohjphemus) and in a snake's burrow, in Texas on 
hogs, and in California. We have a specimen taken at Burnet, Tex. 

In the Canadian Entomologist for 1900, page 20, Lounsbury men- 
tions the possibility of the African species 0. savignyi being intro- 
duced and identical witli 0. turicata. He states, "Neumann in his 
monograph does not give extensive ground for separating 0. savignyi 
and 0. tuiicafa. In this country [South Africa], natives are known to 
carry the tick unintentionally with their belongings from place to 
place. It might easily have been introduced into America with slaves 
in the last century or earlier, just as negroes returning to Africa are 
said to have introduced here the jigger flea (Sarcopsylla penetrans L.). 
This latter insect continues to spread and is now found as far south as 
Durban, Natal." 

Family IXODIDJE. 

The members of the family Ixodidae from their structure naturally 
fall into two subfamilies as suggested by Salmon and Stiles. The 
first, Rhipicephalinip, comprises the forms with short, more or less 
conical palpi, represented m tliis country by the genera Rhipicephalus, 
Margaropus, Ha^maphysalis. and Dermacentor; the second, subfamily 
Ixodinje, includes forms with long palpi and is represented in the 
United States by the genera Ixodes, Amblyomma, and Ceratixodes. 



CLASSIFICATION AND HABITS OF TICKS. 



47 



Subfamily RHIPICEPHALIN^. 
Genus RHIPICEPHALUS. 

The genera Rliipicephalus and Margaropus (formerly Boophilus) 
are structurally sb similar that Neumann and Fuller consider our M. 
annulatus as belonging to the genus Rhipicephalus. All species of 
Rhipicephalus, so far as studied, including five South African forms 
investigated by Lounsbury and one taken by us, drop from the host 
for the purpose of undergoing at least one of the molts. The three 
species placed under Margaropus that have been studied pass both 
molts upon the host. This would seem to supplement the structural 
differences in indicating the generic validity of Margaropus. 




Fig. 3.— Rhipicephalus sp.: Capitulum ol female, ventral view. Greatly enlarged (ciriginal) 



BROWN DOC TICK. 



( lihipircplialiis sp. i 

Salmon and Stiles in 1901 stated that they were not acquainted 
with any North American species, although they called attention to 
the fact that R. sanguineus had been reported by Neumann from 
Panam.a, R. bursa americanus from Jamaica, and an undetermined 
species from Porto Rico. Banks ° records an undetermined species 
from Colorado. Wliat appears to be a new species has been taken 
quite generally from dogs in the southern part of Texas and also 
at Tlahualilo, Durango, Mexico (see figs. 3 and 4 and PI. Ill, fig. 5). 



oThe Arachnida of Colorado. Ann. N. Y. Acad. Sci., Vol. VIII, 1905. 



48 



NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 



It may be called the brown dog tick. Our specimens are from nine 
different localities and were all taken on dogs. Unlike Dermacentor 

variahilis, Mr. Mitchell in- 
forms us, this species occurs 
on all parts of the body of 
dogs. Nathan Banks in- 
forms us that this form is 
closely allied to R. sangui- 
neus. It is probably the 
same as was referred to as 
/?. sanguineus in the annual 
report of the Bureau of Ani- 
mal Industry for 1905, page 
.35. 

Lounsbury has f oimd five 
species belonging to the 
genus Rhipicephalus that 
transmit African coast fever 
in cattle. The possibility 
of the transmission of dis- 
ease by the species we have 
found remains to be inves- 
tigated. TIk^ following are notes we have made on the life history 
of this form: 

Table X. — Oviposition of Rhipicephalus sp.,from dog. 




Fig. i. ^Rhipicephalus sp.: Coxae of 

(ircatly I'lilargrd (original) 



First eggs deposited. 


Oviposition completed. 


Period of Period 

ovi- from 

position, dropping. 




Mays 


Days. Days. 




Mays 


31 37 


Apr. 7 


Mayl 


2S 33 


Apr 8 


Apr. 27 


20 29 




Mly7.::::::::::::::::::::;:::::::::: 


20 1 ,3!) 








Average 


20 i 35 







In the above lot of five ticks collected March 29, as will be seen, the 
maximum period of oviposition was 34 days, the minimum 20, with 
an average of 26. The maximum number of eggs deposited in a lot 
of seven ticks collected July 22 was 1,270, the minimum 91, with an 
average of 636. An engorged tick collected April 30 commenced ovi- 
position May 11, continuing for 12 days, as follows: 

Table XI. — Rate of oviposition in Rhipicephalus sp.,from dog. 



Number of eggs deposited— 


Total. 


May 
10 


„.. 


May 

12. 


f,'.' 


■s.^ 


May May May 
IS. 16. 17. 


May 
18. 


fr 


May' May 
20. j 21. 


May 
22. 


May 
23. 


M.. 


S'' 





73 


179 


235 


181 


107 88 108 


110 


70 


27 20 


9 








(«) 


1.209 



72, Bureau of Entomology, U. S, Dept. of Agriculture. 




The North American Fever Tick and Other Species. 

Fig. l.—M(uv<u-<,i,n^,nnnil,if„:<. male. Fin. ■l.—II:,„,u,,hii.<,ilisl,,,ori<'n'ihi^tris. iVmale. Fig. S.— StiR- 
mal plate of Mnnininiiii^ niiiinhitiis. male. Fit;-. 4.— Mouth parts nf I.i;,:h s ,;„,k-(i. Fig. .">.— Stix- 
mal plate of i;iiii>i<-(jili(ihis sp., male. FiR. 6.— Sti.Kiiuil plate of Aiuhhitmnini inaruhifiiiii. female. 
Figa. 1, 2, mncli eiilarwd: tig. 4, more enlarged: tigs. ;i .5, (i, highly magiiitled. (Original.) 



72, Bureau of Entomology, U, S, Dept of Agriculturi 



2 , 




■m^' 











^ 



fe^ 





Stigmal Plates of Ticks. 

Fis. 1. -Stigmal philfsand anus ui Ih ninir, „i,„- „it, „.<. iumIc. Fifj. 'J.— Stigrmal plate .if same 
Fig. 3.— Stigmal plate ol Anihh/onnua raj, ,nirn.« , rualr. Pig. -1.— Same, female. Fig. •^.— .sii.a 
mal plate of Dermncentur mridhili.--. female. Fig. 6.— Stigmal plate of Bermaceittor occhlciitdlh 
Highly magnified. (Original.) 



CLASSIFICATION AND HABITS OB^ TICKS. 49 

The incubation period of eggs deposited during the middle of April 
was 6 weeks and of those deposited at the end of that month, 33 days; 
seed ticks that emerged the first of June lived for 10 weeks, when kept 
in test tubes on sand. 

Attempts were made to rear this species by placing seed ticks on 
dogs, but with poor success, as few seemed to attach. A dog was 
infested with seed ticks on May 29, but none could be found attached. 
On July 13, however, three adult ticks, two partially and one nearly 
fidly engorged, were found between the toes on a front foot. One of 
the small ticks dropped July 14 and the engorged one on July 15. 
In the middle of October several males of this species appeared upon 
the dog, and these must have come from the above lot of seed ticks. 
They were found to change the location of their attachment from 
day to day. Our conclusion is that this species drops to the ground 
for both molts. 

Genus MARGAROPUS. 

Neumann stands alone in suppressing Margaropus (Boophilus) 
under Rliipicephalus. Although the two groups are closely related, 
it seems evident, not only from their structure but from their habits, 
that they form two distinct genera. 

Neumann's latest arrangement of the forms is as follows: 

R. {M.) annulatus (Say). Southern United States. 

E. {M.) annulatus var. australis (Fuller). Australia, the Antilles, 
and South America. 

R. (M.) annulatus var. calcarata (Birula). North Africa. 

R. (M.) annulatus var. decoloratus (Koch). South Africa. 

R. (M.) annulatus var. caudata Neumann. Japan. 

R. (M.) annulatus Y&T. .argentina Neumann. Province of Buenos 
Ay res. 

Neumann considers that R. (M.) microplus (Canest.) is very prob- 
ably identical with R. (M.) annulatus var. australis. 

The first portion of this bulletin deals with the sole North American 
representative of this genus, M. annulatus (see PL III, figs. 1, 3). 

Genus DERMACENTOR. 

This genus is characterized by the presence of cleft front coxae in 
both sexes, the fourth pair of an immense size in the male but normal 
in the female. The structure of the stigmal plate furnishes valuable 
characters in specific determination in this genus as do the porose 
areas. 

Salmon and Stiles in 1901 had but three species before them from 
the United States, which they identified as D. electus Koch (varia- 
lilis Say), D. reticulatus Neumann, and D. variegatus Neumann. 

5795— No. 72—07 4 



50 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

The species which they then hsted as D. retindatus Neumann is now 
considered by Doctor Stiles and Mr. Banks as Neumann's D. occi- 
dentalis, described from specimens collected in Sonoma County, Cal- 
ifornia, and labeled D. occidentalis by Marx. The D. reticulafus of 
Salmon and Stiles is now considered by Banks as alhipictus Packard. 
While D. reticulatus Fabricius is widely distributed, being found in 
Europe and Asia, so far as known it has not been taken in this coun- 
try. D. parumapertus, described hj Neumann in 1901 from 4 female 
specimens taken at Lakeside, Cal., labeled as taken on a man and in 
a chicken house; and D. hifurcatus Neumann, from a wild cat in 
Texas, described as Ixodes and later referred to the genus Derma- 
centor, seem to come close together, although they may be distinct 
species. According to Mr. Banks, Ixodes nigrolineatus Packard is a 
Dermacentor. To those referred to can now be added Dermacentor 
nitens Neumann, which has been collected by Mr. J. D. Mitchell, of 
this Bureau, making a total of 7 described species so far known to 
occur in the United States. 

AMERICAN DOG <)!{ WOOD TICK. 

[ Dfrnidrenior raridbilis Say.) 

Synonymy (on the authority of Mr. Nathan Bankss): D. mnerivanus authors (not 
L.); D. clcctus Koch, 1844; Ixodes albipictus Pack. (1st Peabody Acad. Rept., p. (iti, 
not Guide and Am. Nat.); /. quinqnestriatus Fitch, 1871; /. robcrtsoni Fitch, 1871 ; /. 
pitnrtiiJiiliis Say. 1S^21C.'), 

This s})ecics is distinguished by the fmely j)unctate stigmal plate 
(see PI. IV, fig. 5) . It is widely distributed over the country, and has 
been taken commonly in northern and southern Texas and in Florida 
on the dog. In some sections of Texas Arnhli/oriinia maculatum and 
RMpicephahis sp. are the common ticks on the dog, which is also the 
case with /. scapularis in Florida. Neumann records a male taken 
on a rabbit, Lepus callotis, by Duges, at Guanajuato, Mexico. Cattle 
also serve as hosts. 

Prof. H. A. Morgan records 7,378 eggs as deposited by a single tick 
between May S and 2(1. These eggs commenced hatching on August 
20, an incubation ]>eriod of 27 days. Our records include data on the 
deposition of eggs by a tick collected April 30, oviposition commenc- 
ing Mav S. The details follow: 





T A B 1. K NIL. Or ipos it ion of Dennac 


■nlor variabilis 










H 


XimilxT of pggs depos 


ited 


- 


c, : 2 ;: :■ i = :: i 1. ' 5 ' 2 ' g 


?S , Si 


s ! ?; , s 


S 


S 


s 




ii 




1 


5" 


i, ' i, 1 




& 

s 


1 


^ 
& 


32 


72 124 232 251 320 237 246 217 257 ^235 253 ,153 

II! 1 1 1 1 ! 1 


35 


104 


34 1 6 , 








(a) 


2802 



CLASSIFICATION AND HABITS OF TICKS. 51 

In five lots of eggs deposited in April the incubation j^eriod varied 
from 37 to 43 days. Four lots deposited during May prior to the 10th 
hatched in 33 or 34 days. Seed ticks that hatched from eggs dejiosited 
May 10 lived until November S or 10, being alive November 6, but 
all were dead on November 10. The i)eriod of survival was thus six 
months from deposition. 

Under summer temperature Prof. II. A. Morgan found engorgement 
of the adult to take place in from 5 to 8 days. He concludes that the 
larva^. and nymphs attach to mammals other than cattle, as the s])ecies 
has only been found on cattle in the adult stage, and attempts to cause 
seed ticks to attach failed. 

np:t tick. 

{Dermacnitnr orcidcntalis Neiuiiaim.) 

This species (see PI. IV, fig. G) was received by Marx from Occi- 
dental, Cal. He determined it as a new species, labeling it D. occi- 
dentalis. Several writers have made use of this name, but it remained 
for Neumann to describe it for the first tune in 1904, placing it as a 
variety of D. reticulatus Fab. Curtice in 1892 referred to it briefly as 
D. americanum (variabilis). It is now considered by Banks to be a 
distinct species and is the one referred to by Salmon and Stiles as 
reticulatus. The true D. reticulatus Fab. is not represented in our col- 
lections, although it may possibly be found to occur here when a 
thorough tick survey is made. 

The species seems to be a western one, being found in the region of 
the Rocky Mountains especially. The Bureau of Entomology and 
Marx collections contain specimens from California, Washington, 
British Columbia, Colorado, New Mexico, and Texas. Salmon and 
vStiles also record specimens from Oklahoma and Tennessee. In a 
specimen taken from a deer skin at Kerrville, Tex., we have what is 
apparently this species, there being some doubt because of the poor 
condition of the individual. The recorded hosts include cattle, 
horse, sheep, deer, and man. 

EI.K TICK. 

( Dermaccntor albijyictns Packard.) 

Dei-maccntor albipictus Packard, Am. Nat., II, p. 559; Guide to the Study of In- 
sects, 9th ed., -p. 662. Not 1st Kept. Peabody Acad., p. 66. (See Banks, A Cata- 
logue of the Acarina or Mites, <rProc. U. S. Nat. Mus., Vol. XXXII, 1907, p. 608.) 

This species is found commonly on the wapiti or ''elk" {Cariacus 
canadensis) in the States of Washington, Montana, Nebraska, Nevada, 
and New York. Salmon and Stiles report that game keepers on the 
reserve of the Blue Mountain Forest Association complain of its being 
very common on the wapiti and that it kills numbers of them. 



52 



NORTH AMERICAN P^EVER TICK AND OTHER SPECIES. 



They suggest the possibiHty of disease being transmitted by it. The 
specimens ^in the Bureau of Entomology collection from Nebraska 
were taken on the beaver. 



TROPICAL HORSE TICK. 

{Dcrinaccnior 7iitnis Neumann.) 

This species is readily distinguished by the characteristic structure 
)f the stigmal plate. (See PI. IV, figs. 1,2, and text figs. 5 and 6.) By 

collections by Mr. 
J. D. Mitchell fi'om 
the ears of horses at 
Brownsville and at 
Harlingen, Tex., this 
species has been 
added to the list of 
ticks found in this 
country. The spe- 
cies was described 
by Neumann in 
1897 from specimens 
in the Marx collec- 
tion, locality un- 
known, and from 
specimens from San 
In lOOl Neumann listed it 




Dermacentor nitrns: Capitulum of fomalo. 
(original). 



Domingo and Jamaica on the horse 

from Guatemala, Venezuela, and Porto Rico 

A single specimen, appar- 
enth^ of this species in the 
njanphal stage, was taken at 
Kerrville, Tex.,byMr. F. C. 
Pratt from a deer skin that 
had been removed in January. 

Genus H^MAPHYSALIS. 

The ticks belonging to this 
genus may be readily recog- 
nized by the presence of con- 
spicuous lateral prolonga- 
tions on the second palpal 
segments. The eyes are ab- 
sent; the coxae of the male 
are all provided with spines, 
those of the female with small 
tubercular processes. Neu- 
mann mentions two species fi'om North America 




Dermacentor nitens: Coxae of male and fi'male. 
Oreatly enlarged (original). 



//. leporis-palustris 



CLASSIFICATION AND HABITS OF TICKS. 53 

and //. concinna, but Banks informs us that he has not found th(> 
latter in tliis country. He recognizes //. cJiordeilis Packard as tlu^ 
form occurring in the eastern United States. 

A South Afi-ican species, //. JeacM, has been determined by Louns- 
bury to transmit mahgnant jaundice of dogs. The larva and nymph 
both drop fi-om the host to molt. Both engorge quickly, sometinu^s 
in less than 48 hours from the time they attach; usually, however, 
remaining from 65 to 75 hours. 



HABHIT TICK. 

{IhevHiphiisdlix 1( pon's-jxilii.^li 
Synonym: Gonixodes rostralis Diu'es. 



Packard). 



Packard described this species (see figs. 7, 8, and PI. Ill, fig. 2) in 
1869 from a female sj)ecimen collected at Fort Macon, N. (\, on a 




Fig. S.—Hscmap,\ y.-alis leporis-palustris: Coxa' of 
male and female. Greatly enlarged (original). 

i-abbit, Lepus palustris. Marx 
reported the species as quite com- 
mon in Kansas, Texas, and Cali- 
fornia. Dr. Cooper Curtice has 
taken an engorged female from a 
horse in Texas, and it has been col- 
lected in Mexico, the host not being given. Neumann mentions an 
engorged female in the museum of Paris labeled ''from Brazil" and a 



Fig. 1 —Uxmaphy sails leporis-paluslris: 
ulum and scutum of female, dorsal 
Greatly enlarged f original). 



54 



NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 



molting n}' mph taken on a Paradoxurus by P'orbes in the Malaysian 
archipelago. Lahille reports it from Argentine Republic. During the 
past two years it has been collected in Texas by agents of the Bureau at 
eight different times from rabbits and hares. The species seems to be 
generally restricted to the genus Lepus, but two instances of other 
hosts having been recorded and these probably accidental. On the 
rabbit the ear seems a favorite place for attachment. As a more 
extensive collection of the tick is made it will undoubtedly be found 
to occur in a much greater territory than is now known. A few notes 
on the life history of this species have been made by us. 





' 


\\KI.J 


; XIII. — V i posit ion 


of Unnidj 


Jnjml 


,s Up 


>ris-p(ilustris. 






Number of eggs deposited— 


Total. 


lo- 


May 1 May 


iviay 


May 
24. 


May 
25. 


May 
26. 


May May 


May 
29. 


May 
30. 


May 
31. 


June 
1. 


June 

2-3. 


2G0 





no 



47 



228 

70 



146 
39 
48 



•1 

43 


98 
7 
29 
17 


72 
3 
22 
22 


63! 4 
: 
14 12 
21 13 




^l\ 
4 











(a) 


1,112 
57 
326 
99 


7 
9 


13 


7 
1 


(a) 



Table XIV. — Period of incubation and longevity of fl:rmaph>isalis leporis-palusliis. 



Eggs depos- Hatching 
ited. commenced. 


Minimum 

inc'ui)a- 

tion period. 


Remarks. 


May 20-23 

May 24-29 

September 2-5 . 


June 20 

June 23 

September 26 . 


3') 
24 


March 5, dead. 
March 5, dead. 
May 11, dead. 



Subfamily IXODIN^. 

Three genera of this subfamily, CVratixod(»s, Ixodes, and Ambly- 
oinina, are represented in the fauna of the United States. They may 
be readily distinguished. The Ixodes have long club-shaped palpi, 
the form of the tliird segment being typical of the genus (see PL III, 
fig. 4). The male and female have only the first coxjb provided with a 
spine of varying size; the anal groove surrounds the anus anteriorly, 
opening posteriorly; eyes are present. In Amblyomma the palpi are 
long but cylindrical; the male has a long spine on the first and fourth 
coxie, while in the female only the first coxa^ have spines, the others 
possessing tubercles; the anal groove surrounds the anus posteriorly 
and opens anteriorly. Ceratixodes, with pointed palpi, occurs on sea 
birds. 

Genus IXODES. 
(The Castor Bean Ticks.) 

Previous to 1796 all ticks were placed under the Linntean genus 
Acarus. In that year Latreille erected the genus Ixodes, giving /. 
ricinus as the type species. For many years following all ticks 



CLASSIFICATION AND HABITS OF TICKS. 55 

were described as belonging to this genus, as is the case with the 
species of Say, Fitch, and Packard. 

Of the genera represented in the United States this has the hirgest 
number of species. Fourteen are recognized, as follows:" 

/. ricinus L., I. frontalis Panzer, I. scapularis Say, I. fuscus Say, 
I. brunneus Koch, /. urise White, /. cookei Packard (wSynonynis: 7. 
cruciarius Fitch, and /. hexagonus of S. & S., on the authority of 
Nathan Banks), /. arcticus Osb., I. diversifossus Neuni., I. dcntatus 
Neum., /. angustus Neum., I. 'mchoatus Neuni., I. sndptus Neuni., and 
I. californicus Banks. 

Of these, four have been taken in Texas by agents of this Bureau — 
ricinus, cool'ei, scapularis, and sculptus. 

Comparatively little is known of the life cycle of the species of 
Ixodes. The work of E. G. Wheler on ''Louping III and the Grass 
Tick" (/. ricinus) is about all that has been done. It seems prob- 
able that all species drop to molt. 

As Mally ascertained to be the case with /. pilosus in South 
Africa, we have found engorged females of I. scapularis to dry up 
in captivity before ovipositing. From Wheler's studies and our 
experience moisture seems to be a necessity in order that molting 
may take place. 

The longevity of the larvae of ticks of this genus must be excep- 
tional, even when some of the long-lived s])ecies of other genera 
are considered. Wheler mentions larva? (7. plumbeus?) which 
hatched on October 9, 1898, from eggs laid in August, that lived 
until the beginning of August, 1899, or about 10 months. They 
were kept in a bottle with a sprig of moss and some damp sand. 

AMERICAN CASTOR BEAN TICK. 

(Lrodcs cookri Packard.) 

Synonymy (on the authority of Natlian Banks) : Ixodes cruciarius Fitch ; /. licragnnus 
S. & S. (not hexagonus Leach, 1815); I. hexagonus var. longispinosa Neuni. 

This species(P]. Ill, fig. 4) was described by Packard from speci- 
mens on a woodchuck, Arctomys monax, at Salem, Mass. Neu- 
mann, in 1899, placed the American form with the European 7. 
hexagonus, but in 1901 he separated the two as varieties. Salmon 
and Stiles, in 1901, followed Neumann, who at that time had all their 
specimens. Banks has examined the National Museum material 
and considers longispinosa the same as 7. cookei of Packard, but 
states that there may be a true hexagonus in this country, though he 
has not seen it. 



a According to Nathan Banks, /. nigrolineatus Packard is a Dermacentor. 



56 NOKTH AMERICAN B'EVER TICK AND OTHER SPECIEi^. 

We have taken this species in Texas from a goat, a raccoon, and 
a skunk. Neumann records it as taken in the United States from 
the otter, mink (Putorius vison), sheep (Texas), spermophile, domestic 
cat (Maine), fox (Cohirado), weasel, porcupine, and marmot. 

EUROPEAX CASTOR BEAX TICK. 

il.rodis rifiuiis LinnaMis. ) 

From its general distnl)Utioii throughout Euroj)e this species has 
been called the European castor bean tick. 

Mr. Banks informs us that in this species the front tarsi are 
longer than in am^ other species of the genus found in the United 
States; that in structure it is very similar to /. scajmlaris, but the 
porose areas are larger and closer together and the scutum is more 
angulate on the sides than in that species. Neumann, in his list 
of hosts of the adult stage, includes sheep, goat, cattle, horse, stag, 
roebuck, dog, cat, fox, ferret, hedgehog, and man; the nymphs and 
larv.ne having occasionally been found on lizards, birds, hares, rab- 
bits, squirrels, ])olecats, ferrets, hedgehogs, mules, bats, and mice. 
In the United States he lists it from Maryland, ''Carolina," Florida, 
California, Kansas, and Texas, on Lepus sylvaticus, Fells pardalls, 
cattle, opossum, gray fox, panther, and wild cat. While we have 
expected to find it frecjuently in Texas, in but a single instance has 
it been taken. This was by Mr. F. C. Pratt, at Mountain Home, 
from a dog. In Louisiana Prof. II. A. Morgan reports it as found 
on mink in all its stages, but on cattle only in the adult stage. 

Although this is an old and widely distributed species, but little 
study has been made of its life history, that of E. G. Wheler, of 
Englantl. in 1S09, standing practically alone. His most valuable 
studies were made to di^termine the relation of the tick to "louping 
ill" of sheep, with whicli.in the light of present knowledge, it seems 
to have only an accidental connection." The following is the sub- 
stance of Wheler's observations ma(h» in England and published in 
1899: 

The a(kut fenuiles are readily recognized before they become dis- 
tended by their deep-red bodies, dark-brown legs, sliield, and other 
points. The males are of a uniform dark brown. A record of 2,050 
eggs fi-om a single female is given, and a very interesting account of 
the remarkable process of oviposition. I^arvae, upon fmding a host, 
attach, and remain for aliout 2 days, by wliich time the}^ are distended, 
black, and globular. Fully distended larva? received February 7 and 
kept in a bottle became hard, dry, and torpid, but on April 29, after 11 
weeks, were found to have changed into nymphs and resumed active 
habits. After molting the nymph takes up its position on the herb- 
age, just as the larvae had done, for a chance of attachment to a host. 

o Journal of the Royal Agricultural Society of England, Vol. X, pt. 4. 



CLASSIFICATION AND HAHITS OK TICKS. 57 

Whereas adults seem to, confine themselves niostly to sheep, cattle, 
and deer, the larvae and nymphs attach themselves very readily to 
various hosts, such as horses, dogs, and human beings. About a 
dozen distended nymphs taken from sheej") May 29, though kept 
moist, had the same dry appearance as the larvae, as before described. 
These molted about July 19, about 11 weeks after removal from 
the host. For some time previous they had appeared dead, no motion 
of the extended legs being perceptil:)le. Of these about half proved 
to be males. About 10 days passed before the sexes attained their 
proper color and strength. On reacliing the adult stage both males 
and females again wait on herbage for a passing host. At tliis time, 
as well as after distention of the female on the host, an action which 
appears to be sexual intercourse freely takes place even in confine- 
ment. The rostrum and other mouth parts of the male are inserted 
in the genital opening of the female, which is situated between 
the bases of the posterior pair of legs. On the host the females grad- 
ually distend, and in the course of so doing vary much in color and 
appearance, so much so in this case that it is difficult to believe that 
they are of the same species. Of the exact periods of engorgement 
we are not informed. Under unfavorable conditions larvae have been 
kept alive for 4 months. NAmiphs were kept alive for 6 months and 
adult males and females for 4 months, being still alive at the time of 
writing (1899). Without moisture, when kept in a dry empty bottle, 
neither larvae, nymphs, nor adults survived more than 2 or 'S chiys. 
Females exposed to 25° F. for a night were found to be but little 
affected. 

This species, according to recent investigations of Kossel and others 
in Germany, transmits European piroplasmosis of cattle. The fact 
that the organism Piroplasma higeminum is the same as is found in 
tliis country lends great interest to investigations to determine 
whether /. ricinus may not transmit Texas fever in the ITnited States. 

BLACK-LEGGED TICK. 

{I.rodcs srapularis Say.) 

Say states that tins species is rather common in forests, and is fre- 
quently found attached to dift'erent animals. Xeumann has not 
recognized the species, but Banks has identified it with a form com- 
mon in some parts of the South. In Texas we have collected it from 
deer and dogs, and in Florida from dogs. In the latter State it seems 
to be very common, and was taken at Havvthorn, Orlando, and Fort 
Myers. 

This species is remarkable for the size that the engorged seed tick 
reaches. These are as large or larger than the engorged nymph of 
Margaropus annulatus, although the adult is not as large as the adult 
Margaropus. Large numbers of seed ticks and adults have been 



58 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

taken on dogs, though as yet not a single nymph. Tliis suggests the 
remote possibiht}^ that the species may pass from the larval to the 
adult stage at a single molt. 

SrrLPTl RED TICK. 

{Ixodes sculptus Xtniinann.) 

This sj^ecies was descril)ed in 1904 from a female specimen taken 
with a female /. ricinus in the Santa Cruz Mountains of California. 
So far as known it had not been taken since until Mr. F. C. Pratt col- 
lected it on prairie dogs at Sherwood, Tex., November 2, 1906. 
More extensive collection will undoubtedly show a wider distribution 
than is now known. 

Genus AMBLYOMMA. 

The species of the genus Amblyomma are distinguished by the 
palpi, which are long and cylindrical. The male has the fn-st and 
fourth coxa^ armed with a long spine; the female has only the first 
cox;e with a spine, the others with tubercles. As far as known all the 
species drop from the host for each molt. Lounsbury has found 
Amhlyomnia lichrnum to be the transmitter of heartwater in cattle, 
sheep, and goats, and has carefully worked out its life history. 

The genus is represented in the Ignited States by four species — 
americanum, cajerniensc, maculaturu, and tuberculatum. A fifth 
species, A. multipunctum, is described by Neumann from two speci- 
mens taken on a tapir and an antelope (Dicrmiocerus furcatus) in 
''North America." These are reported as collected by Donckier. 
As no species of tapir is found north of Nicaragua it seems probable 
that ^'1. ninltipundunt nmst have been tak<ui from that section of 
North America. 

Tabi.k for Si:i'ai!ati\(; tiii-: Si'kciks ok Amhi.vomma of the United States. 
(Adaplt'd from Xciiinann. i 

MALE.S. 

1. Marginal groove exlomling around the posterior Ixirder 2 

Marginal groove not exlending around the posterior l)order .1. liihcrnilulum. 

2. Coxse I bicuspid -^ 

Coxte I armed with one long spine -1- mandatum. 

3. Punctations of scutum lacking from the triangular projections, flat, radiating on 

posterior half -l- rajmnmsc. 

Punctations distrilnited over entire surface of scutum A. ameriranvm. 

FEMALES. 

1. Coxie I bictispid 2 

Coxa? I armed with one very long spine -l- iiiaciilatirin. 

2. Scutum triangular, posterior-lateral borders nearly straight S 

Scutum cordiform, oval, or pentagonal, jiosterior-lateral borders convex. 

A. tuberculatum. 

3. Eyes in front of anterior third of scutum A. cajmnensc. 

Eyes at plane of or behind anterior third of scutum A. americanum. 



CLASSIFICATION AND HABITS OF TICKS. 



59 



LONE STAR TICK. 



(Ambhjomma nmmcanum Linngcus.) 

The lone star tick derives its name from the l)right-silv(>ry spot (m 
the scutum of the female. It is widely distributed, having- been 
reported from Labrador to Florida, and also from Guatemala and 
Brazil. After Margaropus annulatus, it is the most important tick 
in the United States. Though found more commonly on cattle we 
have taken it from man, horse, mule, dog, goat, hog, deer, scjuirrel, 
and wolf, and it appears to attack mammals generally. In portions 
of Louisiana and Texas it becomes a pest of considerable importance 
to moss gatherers and other persons who spend much time in the 
forests. It has been repeatedly taken in Texas during the summer of 
1906 on the clothes or attached to the body of agents of the Bureau. 
Packard mentions a case in which one buried itself in the arm of a 
young girl, producing a raised tumor. 

In May it was found on a herd of dairy cows near Dallas in large 
numbers, though only an occasional specimen of Margaropus annu- 
latus was present. The proprietor of the dairy stated that they were 
very annoying through their attaching to milkers. Mr. J. D. Mitchell 
has found it in the vicinity of Kcrrville and Llano, T(>x., to be the 
most important species, being much more numerous than the fever 
tick. In that region the cattle suffer greatly from it. Its abundance 
seems to be due to the large numbers of sheep and goats kept in that 
section. These serve as hosts, spreading it broadcast, at the same 
time reducing the bunches of Margarojms annulatus seed ticks. 





Table XV. — Ovrposilion of Amblyom 


m,a americanum. 








Collected- 


Oviposition. 


Period of 
ovipo- 
sition. 


Dropping 
to end 
of ovi- 

position. 


Number 
of eggs 




From— 


To- 


Mar. 26 


May 13 

May 25 

May 27 

do 


Apr. 20 
May 18 
May 21 
June 5 
June 7 
June (i 
June 9 
June 5 
June 10 


Daps. 
16 
14 
9 

12 

11 

14 
10 

15 


Days.^ 

22 
20 

20 
19 
•22 
IS 
23 
10 




Apr. 27 




May 2 




May 15 


2 .108 


May 19 


5,040 


Do 


2, 659 


Do 


do.. . 


1,736 


Do 


do 


950 


Do 

Do 


....do... 
....do.. . 


1,510 
1,306 














Total... 


s 


21.2 


19 137 








2. 126 











Our records regarding oviposition, as shown in the above table, are 
from 10 engorged ticks. It will be seen that the maximum number 
of eggs deposited by an individual was 5,040, deposition continuing 
for 12 days; the minimum, 950, with a deposition period of 10 days; 
an average of 2,126 eggs deposited in 12 days. Morgan records as 
many as 6,519 eggs. 



60 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

Table XVI. — Incubation and longevity of Ainblyoinma ainericanuni. 





Com- 
menced 
to hatch. 


Minimum 
incuba- 
tion 
period. 


Seed ticks aU 
dead. 


Period of— 


Eggs depo-sited. 


Oviposi- 

tion 
to death. 


Hatching 
to deal li. 


Apr. 1.5-17 


June 3 


Days. 
50 
42 
42 
35 
33 


Sept. 13 


Days. 
152 


Days. 
103 


Apr. 2.5 


June 5 












Miiv 2,S /iiid 29 

Mav30;ind 31 


July 1 
July 1 


Nov. ea 

Nov. 22 


103 
177 


129+ 
145 



n One or more alive. 

From the above table it will be seen that the incubation period in 
April and May is about 7 weeks. Eggs deposited in the latter part 
of May hatched in 5 weeks. 

Prof. II. A. Morgan has found some specimens to pass the first 

molt on the host, drop- 
ping in about 10 days fol- 
lowing that molt, or just 
previous to the second 
molt. The greater num- 
ber (h-opped in from 4 to 7 
(hiys. The molting of 
the nymph was found to 
last 6 weeks. Engorge- 
ment of the adult in March 
and April occupied from 
7 to 11 days. 

The resistance of the fe- 
male to water seems to be 
similar to that of the fever 
tick. Seven females were 
submerged for 18 hours, 
all becoming active fol- 
lowing removal from the 
water. The next day all 
hours. Onlv one, an unenfrorged 




Fig. 9.-.4mW 



were again submerged for 
specimen, survived. 

In the adult stage both sexes will reattach, as seems pr()ba])le in the 
larval and nymphal stages. 

Mr. Mitchell has observed adults on grass in tlie act of copulation. 

CAYENNE TICK. 

( Anihhjomnta cnjrnnni.^e Fab. ) 

Synonyms: /. crenatum Say (Banks). A. mirtum Kocli (Banks). I. lie r rem: Duge.s 
(Neuni.), .1. sculplus Berlcsc (Noum.i. 

Tliis species was described from Cayenne in 1794, no host being 



CLASSIFICATION AND HAl^ITS OF TICKS. 



fil 



given. The writers' opinion of the specific identity of specimens from 
Venezuela, on cattle, with Texas specimens, on horse and peccary, is 
confirmed by Mr. Banks. The species may be distinguished from 
maculatum readily by the characters given in the table. (See also 
figs. 9 and 10 and PL IV, figs. 3, 4.) 

The species has been reported from, and seems to be generally dis- 
tributed through Mexico and Central America. It has been reported 
from Colombia, -Venezuela, Brazil, and Argentine Republic in South 
America. Neumann also reports it from Cuba and Jamaica. In Texas 
Mr. J. D. Mitchell has taken it in Live Oak County from the peccary 
and horse. Mr. Banks reports specimens from I^ouisiana, Mssouri, 
and Florida. In addition to the hosts mentioned it has been reported 
from toad (Bufo agua), capy- 
bara (Ilydrochxrus capyhara), 
an ant-eater, and man. 

Stoll, in the Biologia Cen- 
trali - Americana, states that 
this species is the most com- 
mon of all Ixodidae in Central 
America, and gives some in- 
formation concerning its hab- 
its. He has never found the 
male in a parasitic state, but 
has found it free on grass antl 
bushes in Guatemala. The fe- 
male, which he states abounds 
in the woods and fields on 
grass and bushes, is occasion- 
ally brushed ofi" by horses, cat- 
tle, or dogs, and even man. 
It adheres tenaciously to the sldn, remaining when undisturbed for 
several days until filled with blood. If forcibly removed, the beak 
breaks ofi^ and remains in the wound, causing a disagreeable and 
sometimes painful inflammation. The young, wliich are distinguished 
by the inhabitants of Guatemala by the name of "mostacilla," hang to 
the grass in clusters of thousands, especially during the dry seasons. 
By their creeping on the skin and frequent biting the^^ form one of the 
greatest plagues of travelers. 

In a letter accompanying specimens of tliis species from Venezuela the 
writers are informed that the ticks do great damage by producing fever 
in cattle, which become weak and in many cases die. It hardly seems 
possible that the malady can be Texas fever; nevertheless tliis species 
may possibly transmit some disease. 




Fig. 10. — Amblyomma cajennenxe: Coxte of 
female. Greatly enlarged (original) 



62 



NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 



Tablk XVII. — Record nf deposition of an engorged tick, Amblyomma cajennense, col- 
lected on peccary April 29. 



Number of eggs deposited— 


67 


o 
1 
85 




1 

278 


2 

1 




S 1^ 22 
213 336 307 


2 S 
1 1 
206' 119 


>> 

60 


1 

68 


1 
33 


1 


1 


ll' 3 







II 
0(a) 


Total 


44 


259 289 


33 


23 


2,650 



a Dead. 

It will be seen that oviposition continued for 20 days from IMay 9, 
a total of 2,650 ejjgs being deposited. Eggs deposited May 14 com- 
menced hatcliing June 21, an incubation period of 38 days. Two 

seed ticks from eggs de- 
posited May 14-25 were 
alive November 6, a pe- 
riod of more than 5 
months from deposition. 
The fact that this pro- 
longetl survival included 
summer months shows 
remarkable vitality in the 
seed ticks of this species. 
Seed ticks of Margaropus 
annulatus seldom survive 
over 3 months in the 
summer. 

GULF COAST TICK. 

(Amblgomnm manilatinn 
Koch.) 

Synonyms : A. tig r in u m 
Koch, A. tenellum Koch, A. ru- 
bripes Koch, A. ovatum Koch, 

A. triste Koch, A. complanatmn Berlese (the prccx-ding on the authorityof Neumann), 

Dermacentor ocddentalis Marx of Niles (Morgan). 

This species was described by Koch in 1844 from ''Carolina," no 
host being given. Prof. H. A. Morgan calls our attention to the fact 
that it was referred to as Dermacentor occidentaUs Marx by Niles.« 

It .seems to be the common species along the Gulf coast of Louisiana 
and Texas. (See figs. 11, 12, 13, and PI. Ill, fig. 6.) It was taken 
by Mr. J. D. Mitchell of this Bureau from cattle, horses, dogs, and 
man in Cameron Parish, La., and Calhoun, Jackson, and Victoria 
counties, Tex. In the Marx collection there is a male specimen 




ll.-Ajnbhiomma maculattim: Mouth parts of fi'inalf. 
Greatly enlarged (original). 



aBul. Va. Agric. Exp. Sta., VII, No. 3, pp. 28, 29, PI. IV. 



CLASSIFICATION AND HABITS OK TICKS. 



63 



It soems (jiiite 
illoctod had been 



recorded as taken from cattle at Memphis, Tenn. 
probable that the animal from which the tick was c 
bred in the Coast sec- 
tion. There are also 
several specimens in 
the Marx collection 
taken in Texas. Neu- 
mann reports it from 
Paraguay, Uruguay, 
Brazil, Mexico, and in 
the Ignited States from 
California, Texas, and 
Tennessee, the latter 
based upon the tick 
before mentioned as 
collected at Memphis. 
He mentions 2 males 
and 1 female as being 
taken on a coleopteron, 
Cercus camj^estris, at 
Buenos Aires, Argen- 
tina. Laliille reports 
it from Argentine Re- 
public, where the fa- 
vorite host is the dog. 
He mentions the fact 
that it is used by the Indians as a leech in certain cases of inflammation. 
The male is especially large, much more so and more elongate than 

either A. cajennense or A. aineri- 
canum. Mr. Mitchell reports that 
while he has observed the sexes in 
close proximity on the animals he 
lias as yet to find them in coitu. He 
lias frequently noticed them to copu- 
late after being removed from the 
animal. However, in Argentina, 
Lahille states that several males 
are usually found attached in the 
immediate vicinity of each female. 
Mr. Mitchell states that on one oc- 
casion he found 7 females clustered 
f on a cow between the ear and the 
horn with no male near. In another 
case he found a cluster of 5 females on the neck of a dog with no male 
in the immediate vicinitv. He has also seen instances where as 




maculalum: 
itly enlarge 




Fu;. 13. — Amblyomma maculalum: Scutui 
fomale. Greatly enlarged (original). 



64 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

many as 5 males were located very close together without any 
females, and other cases on various parts in which both sexes were 
found together. This species is very firmly attached to the host. 
While Rhipiceplialus sp. can be removed easily A. ma culatwm can not 
be removed without evident pain. 

Attempts were made to rear the species upon dogs. On October 
11 seed ticks were placed upon two small dogs. Eight were found 
attached on the 15th. For the first five or six days after attach- 
ment the seed ticks increase slowly in size, remaining of a light color ; 
then in a few hours they seem suddenly to engorge with blood. 
The color of the body becomes purplish and afterwards still darker. 
On the fifth day from placing seed ticks on the dogs 8 were found 
still light in color and 1 dark. On the sLxth day 2 were taken that 
had just detached themselves, leaving 2 dark and 4 light on the host. 
On the seventh day 3 were found to have dropped during the night ; 
2 dropped and the remaining 1 engorged during the day. On Octo- 
ber 19 the eighth tick was found to have dropped during the night, 
making a mininmm period of 5 days and a maximum period of 8 days 
for attachment of larval stage. 

Several of the engorged larvie were kept on moist sand to molt, 
but shriveled and died. 

Th(> incubation period for eggs deposited from the 1st to the 11th 
of Se])t(Miiber varied from 26 to 81 days. Seed ticks from these eggs 
were still alive on March 1, 1907. 

LAND TITRTLE TICK. 

{Arnhlyominn tithirndaliini Marx.) 

This species was described by ]\Iarx in 1894 from specimens col- 
lected by Hubbard at Crescent City, Fla., from the Florida land tur- 
tle, Gopherus (Xerohates) polyphemus. Neumann reports a male 
specimen from Cuba. 

We have received 3 specimens collected in February, 1907, by Dr. 
A. W. :Morrill in Florida on a land turtle (Gopherus sp.), presumably 
the same as the host of the original specimens. 

BIBLIOGRAPHY. 

No attempt has been made to make this bibliography complete. 
In Section A, relating to the cattle tick of North America and the 
problems caused by it, the literature is scattered through many 
experiment station bulletins, veterinary journals, bulletins of the 
Department of Agriculture, and other publications. Only the more 
important are included. In Section B, relating to foreign disease- 



BIBLIOGRAPHY. 65 

transmitting ticks, the literature is found for the most part in the 
governmental publications of South Africa and Australia, although 
recently important memoirs have been issued in Germany and France. 
As in the preceding section, we give only the more important publi- 
cations that are now available. For a very complete bibliography 
of this subject down to 1903, containing 221 titles, see A. Schmidt 
below. In Section C, relating to ticks as transmitters of human 
diseases, there is at present but a scanty literature. It is practi- 
cally all referred to by us. In Section D, relating more particularly 
to the classification of ticks, practically all of the works dealing 
with North American species are listed, together with the more impor- 
tant foreign publications. For a more extended list, see Salmon and 
Stiles, 1901, below. 

SECTION A. 
Relating to the North American Fever-Tick Problem. 

Butler, Tait. 

1902. — The breeds of V)eef cattle and l)eef ])n)ducti()ii in Norlli Carolina N. C. 
State Bd. Agi-., bul. 23, no. 7. 

Pages 48-51 deal with: "Cattle tick an obstacle to tlic dcvclopincnt of the cattle 
industry." 

1903.— Ileport of State Veterinarian in Rept. Comm. Agr. N. ('. for 1902, pp. 40-47. 
Regarding the cattle quarantine, the extermination of the cattle tick, losses from 
tick fever, and the tick an obstacle to the improvement of the quality of the cattle 
of the State. 

1903.— The cattle tick and its relation to tlie cattle industry of North Cai-olina 

<N. (\ State Bd. Agr., bul. 24, no. 5. pp. 28, figs. 2. 
1903. — The cattle tick and the quarantine restrictions <N. ('. Stale Bd. Agr., 

bul. 24, no. 10, pp. 30-37. 
190(i. — Progress made in exterminating the fever tick (Boophiliis annulatns) in 
North Carolina <N. C. Dept. of Agr. circular (unnumbered) issued 
January 1, 1906, pp. 4. Reprinted in part in Farm and Ranch, vol. 25, 
no. 15, p. 7, April 14. 

This circular deals with the most successful work that haslieen done in the practical 
eradication of the cattle tick. 
CONNAW.W; J. \V. 

1897. — Texas fever or acclimation fever <Mo. Agr. Exp. Sta., bul. 37, pp. 81-139, 
figs. 11, April. 
CoNNAWAY, J. W., and M.' Francis. 

1899.— Texas fever <Mo. Agr. Exp. Sta., bul. 48, pp. 66, figs. 11. 
Curtice, Cooper. 

1891.— The biology of the cattle tick <Journ. Comp. Med. and Vet. Arch., vol. 12, 
no. 7, pp. 313-319, July. 

1892. — About cattle ticks Journ. Comp. Med. and Vet. Arch.. January. 

This and the preceding paper were issued together with nidependent continuous 
pagination. 
1892.— The cattle tick <Texas Agr. Exp. Sta., bul. 24, pi). 237-252. 2 pis., 
December. Appendix on preventive measvires by M. Francis. .\1)- 
stract, Insect Life, vol. 5, p. 294. 

This bulletin gave the first details of the life history of the cattle tick. It was a 
pioneer work the value of which has been appreciated by all subsequent workers. 
5795-No. 72—07 5 



6n NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

Curtice, Cooper — Continued. 

1896.— On the extermination of the cattle tick and the disease spread l)y it 
• Journ. Comp. Med. and Vet. Arch., vol. 17, pp. 649-655. 

On page 655 Doctor Curtice gives the first reference we have found to the possibihty 
of totally exterminatingthecattle tick from the United States. " I look most eagerly 
for the cleansing of even a certain portion of the infected territory under the direct 
intention of man, for it opens the way to pushing the ticks back to the Spanish isles 
and Mexico, and liberating cattle from disease and pests and the farmer from untold 
money losses. Let your war cry be: Death to the ticks!" 

1897.— The cattle tick and what may be done to prevent it <The Southern 
Planter, vol. 58, no. 1, pp. 24-27, January. 

This paper contains the original suggestion of the pasture eradication and feed- 
lot systems of tick eradication. 

1897.— Texas fever The Southern Planter, veil. 58, no. 3, pp. 116-117, March. 

1899. — Cattle quarantine, ticks and distemper <N. C. State Bd. Agr., bul. 
(imnumbered), July 1. 

1899.— Cattle quarantine line <N. C. Agr. Exp. Stu., sj)ec. bul. 52, pp. 1-28. 

1905._The cattle-tick plague <The Southern Planter, vol. ()(>, no. 5, pp. .376-378, 
May. 

.\ report of progrcs.s in cxtenniiialion iiiul ii continuation of the two articles pub- 
lished in the above Journal October, l.S9(i, and January. IS'.tT. 

D.\LiiYMPLE, W. II.. II. .\. M<)U(iA\, and W. R. Dodson. 

1898._Cattle tick and Texas fever <La. Agr. Exj). Sta., l)ul. 51, j)]). 230-282, 
figs. 2, pis. i-vii. 

Detailed account of niiiny experiments relating to the life history of Margaropus 
(innuhitii.i. Includes studies of the effects of water, heat, cold, and light on the eggs, 
seed ticks, and adults, and much information on pasture eradication. The plates 
a re excellent. This publication adds gn>a tly to the data furnished by Doctor Curtice 
in Texas Bulletin 24. 

1899. — Immunization against Texas fever l)y Idood inoculaticm <La. Agr. Exp. 
Sta., bul. 57,^ser. 2, pp. 146-185. pis. 2. chts. 3, figs. 3. 

DiNWIDDlE. R. K. 

1892.— Parasitism of donu'slicated animals < Ark. .\gr. Ex]). Sta.,l)ul. 20, pp. 14-31. 

Treats of mites, cattle ticks, etc. 
189(i. — Texas cattle fever in various localities <Ark. Agr. Exp. Sta.. report. 1896, 
pp. 36^0. 

A reprint from bulletin 40 of the .station. 
1898. — Methods of combating communicable diseases of farm animals <Ark. 
Agr. Exp. Sta.. bul. 51, pp. 35-46, May. 

Francis, M. 

1889.— Report of the veterinarian. Tex. Agr. Exp. Sta., second annual report, 
1889, pp. 5&-60. 

.V brief account of the southern cattle plague and its treatment, with notes on joint 
experimentii by the Texas and Missouri Stations. 
1892. — The cattle tick, preventive measures for farm and range; use <Tex. Agr, 
Exp. Sta., bul. 24, pp. 253-256, fig. 1, December. 

1894.— Veterinary Science <Tex. Agr. Exp. Sta., bid. 30, pp. 436-458, March. 

Treats of animal parasites, device for destroying ticks, and influence of ticks in the 
dissemination of Texas cattle fever. 

Francis, M., and J. W. Conxawav. 

1899. —Texas fever <Tex. Agr. Exp. Sta.. bul. 53, p]). 53-106, figs. 13. 

Lewis, L. L. 

1897.— Texas fever <Okla. Agr. Exp. Sta., bul. 27, pp. 8-15, fig. 1, June. 



BlBLIOaRAPHY. (>7 

Lewis, L. L. — Continued. 

1899. — Texas fever. An ai'count of experiments <()kla. Asji-. Kx]i. Sia.. Iml. :','.!, 

pp. 28. figs. 5, May. 
1900. — Means of preventing Texas fever <Okla. Agr. Exp. iSta., report, 1900, 
pp. 26-28. 
Mayer, Au(iUST. 

1906.— The cattle tick in its relation to .southern agriculture <!'. S. Depi. .\gric., 
farmers' bul. No. 261, pj). 1-22. 

.VnexcellpntossMvoii the l.roail aspects of t lie tick problem. 

Mayo, N. 8. 

1897.— Texas fever <Kaiis. Agr. Exp. Sla., huh 69, pp. 121- l:'.4, .Ttiuc 

Mct'ULLOCH, C. 

1899. — The prevention of Texas cattle fever and the amended laws cnnt rolling 
contagious and infectious diseases <Va. .\gr. Ex]). Sl:i., bul. 104, 
pp. 167-180, September. 
Melvin, a. D. 

1906.— How to get ri<l of cattle ticks <U. S. D. A., Bur. A. 1., cir. 97, pp. 1-4, 
1 pi. 

Deals with the haiui-|)icking and greasing, the tie-rope or picketing, jimi the two- 
field methods. 

Miller. W. McC N. 

1895.— Texas cattle fever <Nev. Agr. Exp. Sla.. bul. :^1. i)p. 11, Deccmbet . 

MoHLEK, J. R. 

1906. — Texas fever (otherwise known as tick fever, si)le:ielic fever, or south- 
ern cattle fever), with methods for its prevention <U. S. J). A., Bur. 
A. I., bul. 78, pp. 1-48, pis. 3, figs. 3. 

A comprehensive treatment of the whole subject, including syni|it()ins, p,-i.thology, 
therapeutics, and methods of tick eradication. 
I90(i. — Texas or tick fever and its prevention <!'. S. 1). A., farmers' bul. 258, 
pp. 1-46, figs. 1-6. 
This is a somewhat condensed re-(;dition of the preceding. 

Morgan, H. A. 

1899.— Ticks and Texas fever <La. Agr. Exp. Sta., bid. 56, pp. 128-141, pis. 9, 
September. 

This is the only .Vmeriean work on the life history of species other than Margaroiius 
annulatus. It deals with A mblyomma amencanum , DrriiKui ntor itirinhih.s nlfclus), 
both of which were found to l)e nonpathogenic as far as splenetic fever is <'oncerned, 
and Ixodes ricinus. Many details of life history are given. The plates are most 
valuable. Marx's paper on the classification of the Ixodidse is reprinted as a part of 
this bulletin. 
The starving-out method of eradication is again emphasized. (See Dalr.vmple, 
, Morgan, and Dodson, 1898.) 

1903. — How can we exterminate the cattle tick? <Proc. La. State Agr. Soc. and 
Stockbreeders' Assn.. 1903, pp. 77-79. 
Emphasizes the necessity for cooperation among the fanners. 
1903.— The cattle-tick situation <Proc. Soc. Prom. Agr. Sci.. 1903, pp. 72-74. 

.Votes are given on the lif(^ history- of the cattle tick. 
1905.— Texas fever cattle tick: pasture methods of eradic'ation <Bid. of Agr. 
Exp. Sta. Univ. Tenn., vol. 18, no. 1, pp. 1-10, ligs. 3. 
.\lso published as bul. 82 (second series), La. Agr. Exp. Sta., pp. 1-1.5. 
This publication urges the feasibility of the practical application of a knowledge 
of the life history of the cattle tick in feed-lot and pasture-eradication systems of 
eradication and centered public attention. It gives full details as to procedure. 
See also Dalrymple, Morgan, and Dodson. 



68 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

Newell, Wilmon, and Dougherty, M. S. 

1906.— The cattle tick. Studies of the cK't,' and seed-tick stages. A simple 

method of eradicating the tick <La. Crop Pest Comm., cir. 10, pp. 1-32, , 
hgs. 1-S. 

Contains records of very careful work on the egg and seed-tick stages. The prac- 
tical importance of the work is well demonstrated. 

NiLEs, E. p. 

1897.— The cattle tick in Virginia <Va. Agr. Exp. Sta., bul. 7(i, pp. 45-50, 

May. Southern Planter, July, 1898, pp. 32()-327. 
1898.— A preliminary study of ticks <Va. Agr. Exp. Sta.. Inil. 86. i)p. 25-30, 

4 {)ls., March. 

NoRG.\AKD, VkTOK A. 

1898.— Cattle dipping, experimental and i)ractical V . S. I) A., yearbook. 1898, 
pp. 453-172. 

E.xperiments, principally with saturated solutions of sulphur in <-.\tra dynamo oil 

Ransom, B. H. 

1906.— Some unusual liosi relations of the Texas fever tick , U. S. D. A., 
Bur. A. I., cir. 98, pp. 1-8. 

Details experiments in attempting to cause Margaropus annulatus to develop on 
horses, mules, as.scs, rabbits, dogs, and cats. On the last a specimen developed to 
the adult stage, but did not engorge. Shows that under certain conditions the cattle 
tick can reattach after being artificially removed. Refers to recent European investi- 
gations that show that liodts ricinus transmits European piroplasmosis of cattle, 
and suggests the possibility that yiargaropus unnulat-jx may transmit its disea.se in 
the nymphal or adult states. 

Redding, R. J. 

1889.— Cattle ticks and Texas fever : Ga. Agr. Exp. Sta., bul. 49, i)p. 228-229. 

Riley, C. V., and L. O. Howaud. 

1899.— The Texas c'attle tick • Insect Life, vol. 2. July, 1889, p. 20. Habits 
and remedies. 

Robert, J. C. 

1897.— Acclimation fever, or Texas fever - Miss. .\gr. Ex]). Sta.. l)ul. 42, pp. 32, 

figs. 4. 
1901.— Texas fever <Miss. Agr. Exp. Sta., luil. (I!*, p]). 1 15, tigs. 4, November. 
1902.— Tick fever or murrain in Soutliern cattle (commonly termed Texas fever) 
Miss. .\gr. Exj). Sta.. l)ta. 73, pp. 1-24. figs. 2. July. 

Salmon, D. E. 

1884.— (Geographical distribution of Southern cattle fever In report of the 
Chief of the Bureau of Animal Industry. I^ S. Comm. Agr., report, 
. 1884, i)p. 252-258, pis. 3. 

Di.scusses occurrence of fever in Va., N. ('•, S. ('., Ga., Tenn, Maps show limits 
east but not west of Mississippi River. 
1904.— Relations of Eederal Government to control of c(mtagious diseases of 
animals V. S. I). A., yearbook, 1903, pp. 491-506; pp. 505 et seq. 
Deals with M rgarnpiis i nnul tun. 

Salmon, 1). E., and Theobald Smith. 

1892.— Southern cattle fever (Texas fever) U. S. D. A., Bur. A. I. special 

report on diseases of cattle and cattle feeding, pp. 428^38, pis. 42-44. 
1904. — Texas fever, or Southern cattle fever <U. S. D. A.. Bur. A. I.; cir. 69. 

pp. 1-13. (Reprint from special report diseases of cattle, revised, 1904, 

by Salmon and Mohler.) 



BIliT.TOdKAPHY. 69 

Salmon, D. E., and (". \V. Stilks. 

1901.— The cattle ticks ( Ixo.loidea) <if Uic Inilcd Slates U. S. J). A., lUir. A. 1., 
17th ann. report, 1901, pp. :?80-491, pis. 74-98, figs. 47-238. 

This is a most vahitil)le work, particularly rich in l)iI)liographical references and 
illustrations. It is an absohito essential in the study of the ticks of this country. 

SCHROEDER, E. C. 

1900.— A note on the vitality <>f tlie Sonthern cattle tick <U. S. I). A., B\ir. A. I., 
Ifith ann. report, pp. 41—12. 
ScHROEDEK, E. ('., and W. E. Cotton. 

1900. — (rrowing iioninfected ticks and afterwards infecting tliem <U. S. D. A., 
Bur. A. I.. Kith ann. report, pp. 33-41. 
Smith, Theobald, and F. L. Kilborne. 

1893. — Investigations into the nature, causation, and prevention of Texas or 
Southern cattle fever <U. S. D. A., Bur. A. T., hul. 1, ])]). 1-301, 
pis. 10, figs. 7. 

This scholarly work demonstrated the transmission of fever liy Manjaropu.s annu- 
tatus. It suggests much of the work since done in the study of disease transmission 
by insects and ticks. 

Smith, T., F. L. Kilborne and E. <'. Schroedek. 

1893.— Additional observations on Texas cattle fever <II. S. 1). A., Bur. A. L, 
l)ul. 3, pp. ri7-72. 
Vincenhellek, W. G. 

190(5.- The cattk' tick in Washington and Benton counties< Ark. Agr. Exp. Sta., 
bul. 90. pp. 131-141. 
Willou(;hby, C. L. 

1904. — Cattle ticks and Texas fever; immunizing experiments in (ieorgia<Ga. 
Agr. Exp. Sta., bul. 64, pp. 143-182, figs. 9, August. 

SECTION B. 

Relating to Forekjn Disease-Transmitting Ticks. 
Banks, C. S. 

1904. — The Australian tick (Boopkilus australis Fuller) in the Philippine Islands 
U. S. D. War, Bur. Govt. Laboratories, bul. 14 (Biological Laboratory, 
Div. Ent., bul. 2), pp. 1.3-21, figs. 10-23, pis. 9. 
Bruce, D. 

1905. — The advance in our knowledge of the causation and methods of prevention 
of stock diseases in South Africa during the last ten years <Science, 
n. s., vol. 22, pp. 289-299 and 327-333. 
Edington, A. 

1904. — Note on the co-relation of several diseases occurring among animals in 
South Africa <Agr. Journal Cape Good Hope, vol. 25, p]>. 139-152. 

Froggatt, Walter W. 

1901. — The fowl tick {Argas americanus Packard) <Agr. Gazette N. S. Wales, 
vol. 12. pp. 1349-1352, pi. 

Fuller, Claude. 

1890.- Ticks, a fowl-infesting tick ( Argas ^p.) <Agr. Gazette N. S. Wales, vol. 7, 

pp. 590-597 (reprint, pp. 1-8). 
1896.— The bovine tick fever <Agr. Gazette N. S. Wales, vol. 7. pp. 760-787, 

pis. 1-5. 
1899. — The common blue tick <Agr. Journal Cape Good Hope, vol. 14, pp. 363- 

369, March. 



70 NORTH AMERICAN FEVER TICK AND OTHER SPECIES, 

Gray, C. E., and W. Robertson. 

1902. — Report upon Texas fever cv redwaler in Rhodesia < Argus Printing 

and Pub. Co., Ltd., Cape Town, pp. 27, pis. 22. 
1902. — Redwater in Rhodesia <Agr. Journal Cape Good Hope, vol. 21, pp. 435- 
458, November. 
Herrera, a. L. 

1905. — Las Pardsitos del Ganado <Coni. Parasit. Agrieola., eir. 8, pp. 22-25, 
ligs. 81-34. 
Hunt, Sidney. 

1898.— Tiek fever <Queensland Agr. Journal, pp. 235-23G. 
HUTCHEON, D. 

1900.— Redwater and its history <Agr. Journal Cape (iood no])e, vol. 17, pp. 

331-339, 395-409. 
1900.- History of heartwater <Agr. Journal Cape Good Hope, vol. 17, i)p. 410- 

417. 
1903.— Vindent redwater in llie Transvaal <Agr. Journal Cajx' Good Hope, 
vol. 23, no. 1, })]). 39 (iO. 
JortuN(i, J. W., and P. G. Wooi,i,i:y. 

1904.— Texas fever in the Philipi)ine Islands and the Far East • U. S. D. War, 
Bur. Govt. Laboratories, l)ul. 14, i)p. 5-11, ])ls. 15. 
KossEL, li., A. Weber, ScHiJTZ, and Miessner. 

1903. — Ueber die Hiimoglobinurie der Rinder in Deutschland <Arb. K. 
Gesundheitsamte, no. 1, i)p. 1-77, pis. 3. 

The blood parasite is thf same as thatof ■I'cxas fever. Ixodes ricinus (rcdurius) 
aets as a t ransmitter. 

L.\HILLE, F. 

1905. — Contribution a I'etude des Ixodides de la Repulilitpie Argentine, pp. 
l-l(i(), pis. 1-13. 

This paper t'oiitairis a great lie.-il on tlii' l)i()l(>K.v of Maryaropnx inicropiu.s. 
LiGNIERES, J. 

1900. — La Triste/.a on Malaria bovine dans la Republicjue Argentine, pp. 1-172, 
pis. 14. 

LOUNSBI^RY, C. P. 

1899. — The bont tick Amblyomma hchrpcMm Koeh <Agr. Journal Cape Good 

Hope, vol. 15, pp. 728-743. 
1900.— Life history of a tick <Ent. News, vol. 11. ]))). 33(1-340, Ja!iuary. 

Life history of A mblyomma hebrsfum Koeh. 
1900. — Tick-heartwater experiment <A,gr. Journal Ca})e (iood Hope, vol. 16, 

pp. 682-(;S7. 
1900.— Insect bites and the effects thereof <Can. Ent*., vol. 32, ])p. 17-24. 

Argas and Ornithodoros spp. 
1900.— Notes on some South African ticks <U. J^. I). A., I^ur. Ent., l)ul. 26, 

n. s., pp. 41-49. 
1902. — Report of Government Entomologist for the Cape of Good Hojx' for 1901. 

Ineludes "Tieli heartwater investigations." pp. 29-73, pis. 4-»). 
1903.— The fowl tick. Studies on its life cycle and haliits <Rept. Agr. no. 20, 

pp. 1-15, pis. 3. Reprint Agr. Journal Cape (lood Hope, September. 
1903. — Report of the Government Entomologist for the Ca])e of Good Hope for 

1902. 
"Ticks and Ilhodesian cattlc> di.sease." "Ticks and malignant, jaundice, ricks 

and heartwater," pp. 16-41. 
1903. — Ticks and African coast fever <Transvaal Agr. Journal, vol. 2, no. 5, 

pp. 4-13. 



BTBLIOOmAPHY. 71 

LouNSBURY, C. P. — Continued. 

1904. — External parasites of fowls <Reprinte(l from (lie .V^r. Journal, ])[). 7, 
November. 
Relating to Argas persicux. 
1904. — Persian sheep and heartwater <Agr. Jnuriiul ('a])c Good lIoi)c. vol. iT), 

no. 2, pp. 175-18(), figs. 8. 
1904.^Distribut,ion of coast fever ticks <A<;r. Jounial Cape (iond Hope. vol. 25, 
no. 3, pp. 268-270, pi. 1. 

The distribution of lihipici jiIdiIuk ,i i>pnuUruhiliix. ]:. ftimus, R. crrrlsi. and R. 
capensis is briefly outlined. 
1904.— Transmission of African coasi fcvci- <Kci)l. Agr., no. ■">. pi>. 1-7, j)ls. :5. 

Reprint: Agr. Journal. ('a])c (iood IIoi)c, .\pril. 
1904.— Report of the (iovcrinuenl Kiitomolot^nst for the Cape of (iood llo])e for 
1903. 

Includes "ticks and malignant jaundice" and "ticks and lieartwater," "ticks and 
South African coast fever," pp. 11-45, pis. 7 
1905.— Tests of substances for tick destruction <.'\gr. Journal Cape (iood Hope, 

vol. 2(i, ])p. 387-395, March. 
1905.— Report of the Government Entomologist for the Half Year ended June 30, 
1904. 
Contain.sa special report, "licks and Africancoast fever." pi). 1()--J.">. 

Mally, C. W. 

1904.— Notes on the so-called paralysis tick. Lodes pUo.vis <Agr. Journal Cape 
Oiod Hope, September. Reprint l)y Dc^it . of Agric, no. 17, 1904. 

Marchaux, E., and A. Salimbeni. 

1903.— La Spirillose des Paules <.\nnals 1' Instilul Past<-ur, vol. 17, pp. 5(;i>-5S0. 
Spirillosis of chickens and other I'dwis t nmsiiiif teil by A t-.w.s- inini(iln.'<. 
MOTAS. 

1903.— The role of ticks in llie deveh.innent of carccag <Compt. Rond. Soc. 
Biol., Paris, vol. 55, no. 14, pp. 501-504. 

The writers liMve seen only ,•! review of this paiicr in Kxperinient SIntion Record. 

Pound, C. J. 

1899. — Notes on the cattle tick. Its development, life history. hal)its, and geo- 
graphical distribution <P. Soc. Queensland, vol. 14. ]))). 28-38. 

Robertson, F. H. 

1905.— Fowl tick ex])erimenls <Journ. Dept . .\gr. West Australia, vol. 12. n... (i, 
l)'p. 5(il-5(i3. 

It was found that fowl ticks remain alive at least 2 years and 3 months vvitliout the 
presence of any fowls from which to derive nourishment. In these experiments the 
ticks were kept in small pill boxes which were practically air tight. In the nymph 
stage ticks may live for 2 months without food. 
:^CHMIUT, A. 

1904.— The tick disease of cattle (hcemoglobinamia ixodioplasmatica houni) in 
German and English East Africa and Uganda <.\rch. Wiss. ii. Prakt. 
Tierh., vol. 30, nos. 1-2, pp. 42-101. 

The literature of this subject is discussed with references to a bibliography of 221 
titles. We have not the original work at hand and refer to the translated title in 
the Experiment Station Record. XVI, p. 201. 

Stockman, Stewart. 

1904.— Rhodesian redwatcr, vol Fast African coast fever, vel tropical piroplas- 
mosis <Rcporl of ihc Tran.^vaal Dept. of .\gric.. 1903 to 1904, pp. 
40-66. 

Includes history of invasion and spread in Transvaal, permit- system, dipping, 
immunity, preventive inoculation, prevention and eradication, and transport expert- 



72 NORTH AMERICAN FEVER TICK AND OTHER SPECIES. 

Theiler, a. 

1903 and 1904. — The Rhudesian tick fever < Transvaal Agr. Journ., vol. 1 

(1903). no. 4, pp. 93-110, pi. ] ; vol. 2 (1904). no. 7, pp. 421-438, pi. 1. 
1904. — The transmission of East Coast fever by ticks <Transvaal Agr. Journ., 

vol. 3. no. 9, pp. 71-86, October. 
1905. — Further notes on piroplasmosis of the horse, mule, and donkey <Trans- 

vaal Agr. Journ., vol. 3, no. 12, pp. 706-716. 
1906. — Transmission and inoculability of spirillosis in cattle <Transvaal Dept . 

Agr., ann. rept. Dir. Agr. 1904-1905, pp. 123-151. 
The writer shows that the natural transmission of spirillosis is by the progeny of 

Rhipicephnlus decoloratus, which have developed on cattle affected by spirillosis. 

Wheler, E. (1. 

1899.— Louping ill and the gra.ss tick <J(mrn. Royal Agr. Soc. England, ser. 3, 
vol. 10, pt. 4, pp. (126-644. 
See note under following title. 

1902. — Parasitically inoculated diseases <Trans. Highland and Agr. 8oc. Scol- 
land, ser. 5, vol. 14, pp. 16-35, figs. 2. 

Sunnisesthat "loupingili" is transmitted by Ixodes ricinus. Later investigatitms 
have negatived this. 

SECTION C. 

RKI,ATIN(i TO Tr.WSMI.S.SION OK HUMAN DISEASE I)V TiCKS. 

Christy, Cuthbert. 

1903. — "Tick fever" in man <The Thompson Yates and Johnson Laboratories 
Report, vol. 5, ii. s.. ])art I. pp. 1S7-189. 

DuTTON, J. E., and J. L. Todd. 

1905. — The nature of human tick fever in the eastern part of the Congo Free State, 
with notes on the distribution and bionomics of ilie tick <Liver])ool 
School of Tropical Medicine, memoir 17, pp. 26. 

Includes paper by Robert Nowstead. 'On the external anatomy of Ornithodoros 
mouhnta (Miirrayl." 

King, W. W. 

1906. — Experimental transnussion of Rocky Mountain spotted fever t)y means of 
the tick <l'. S. T. I)., Pul)lic Health and Marine-Hospital Service, 
Puldic Health Reports, vol, 21, pp, 863-864. July 27. 

Newstead, R. 

1905. — On the extenud anatomy of rnithodoros moubata <Liverj)o()l School of 
Tropicid Medicine, memoir 17, pp. 21-26, November. 

Rh'ketts, H. T. 

1906. — The study of 'Rocky Mountain spotted fever"" (tick fever?) by means 
of animal inoculations <Journ. Am. Med. Assn., vol. 47, \)\). 33-36, 
July 7. 
1906. — The transmission of Rocky Mountain spotted fever l)y the bite of the 
wood tick { Dcniiacfntor ocndcntalia) <Journ. Am. Med. As.'<n..vol. 47, 
]). 35S, August 1. 

Stiles, Ch. \V. 

1905. — .V zoological investigation into the cau.se, transmission, and source of Rocky 
Mountain •'spotted fever" <L'. S. T. D., Public Health and Marine- 
Hospital Service, Hygienic Laboratory, bul. 20, pp. 1-121. 



BIBLIOGRAPHY. 73 

SECTION D. 

RELATIN(i TO THE CLASSIFICATION AM) 1 )lSTI{inUTION OF TiCKS. 

Banks, Nathan. 

1895.— The Araclmida of Colorado Ann. N. Y. Acad. Sci., vol. S, j)]). 417-434. 

Dermacentor ameriranu.t L. and Khipitephalus sp. are listerl. 
1899. — Reports upon the insects, spiders, mites, and myriapods collecte<l on the 
(bmmander Islands Expedition (The fur seals and fur-soal islaiuls of 
the North Pacific Ocean, pt. 4, pp. 328-351). 

Lists Ixodes borealis, supposes I fimbriatu.i Kramer and Neumann to he tln' male. 
1901. — Acarina in "Some spiders and other arachnida from southern Arizona" 
<Proc. Y. S. Nat. Museum, vol. 23, p. 590. 

Mentions Argax sanchezi Duges from New Mexico and Arizona; also a species 
of Ixodes from Arizona. 

1902.— Papers from the Hopkins-Stanford Galapagos ExjxHlilion, 1898-1899, 
vol. 7, Entomological Results ((i). The Arachnida, Proc. Wash. Acad. 
Sci., vol. 4, p. 70, pi. 2, fig. 9. 
Argaa tranmersa, n. sp., and mention of A mblyomma pilosum Ncum. 
1902. — Some Arachnida from New Mexico <Proc. Acad. Nat. Sci. Phila., 1901, 
pp. 568-590. 

\Asts. Argas .saiuhtzi Nvuui., Ixodes diversifo.ssus Ncnut.. Drrmmentor retiiiilatus 
Fab., Margaropus (Boophilus) annulatus. 

1904.— The Arachnida of Florida <Proc. Acad. Nat. Sci. Phila., pp. 120-147. 

Margaropus {Boophilus) annulatus Say, Ixodes scapularis Say, Dermacentor 
lariabilis Say, Amblyomma tuberculatum Marx. 

1904. — Some Arachnida from California <Proc. Cal. Acad. Sci., vol. 3, ser. 3, 

no. 13, pp. 331-369, pis. 38-41. 
1904. — A treatise on the Acarina or mites <Proc. U. S. Nat. Museum, vol. 28, 

I)p. 1-114. 

BiRULA, A. 

1895. — Ixodidai novae vel parum cognitse Musei Zoologici Academise Csesareae 
Scientiarum Petropolitana^ <Bull. Acad. Imp. Sci. St. Petersbourg, ser. 
5, vol. 2, no. 4, pp. 353-364, pis. 1-2. 
F^TCH, Asa. 

1872. — Fourteenth report on the noxious, beneficial, and other insects yf the 
State of New York, pp. 363-373. 

Contains descriptions of the following species: Ixodes ame.ricanus, quinquestriatus, 
robertsoni, cruciarius, Ixodes (?) odontalgix. 

Hassall, Albert. 

1900. — Note on the chicken tick {Argas amcricanus) <CU. S. D. A., Bur. A. I., 16th 
report, pp. 496-500, figs. 16-22, pi. 16. 
Koch, C. L. 

1847. — Uebersicht des Arachnidensystems, vol. IV, pp. 1-136, pis. 1-30. 
Marx, George. 

1893. — Note on the classification of the Ixodidte <Proc. Ent. Soc. Wash., vol. 2, 
pp. 232-237. 
Contains tables of North American genera. 
1893. — r)n the morphology of the ticks <Proc. Ent. Soc. Wash., vol. 2, pp. 

271-287. 
1894. — Plate illustrating following species published in connection with obituary, 
Proc. Ent. Soc. Wash., vol. 3, pp. 195-201: Rhynchopriuvi spinosum, 
{=Ornithodoros megnini, nymph), Omithodoros americanus, (=0. meg- 
nini, adult ) , Argas americanus. 



74 NORTH AMERICA N FEVER TICK AND OTHER SPECIES. 

Murray, Andrew. 

1877. — Economic entomology, Aptera, pj). 180-204. 
Deals with the Txodidap. 

Neumann, L. G. 

1896.- Revision de la famille des Ixodides, I <Mem. Soc. Zool. France, IX, 

pp. 1-44, figs. 1-36. 
1897. — Revision de la famille des Ixodides, II. Ixodintp -Mem. Soc. Zool. 

France, vol. 10, pp. 324-420, figs. 1-45. 
1899. — Revision de la famille des Ixodides, III Mem. Soc. Zool. France, vol. 

12. pp. 107-294, 63 figs, in text. 
1901. — Revision de la famille des Ixodides, IV Mem. S()<'. Zool, France, vol. 

14. pp. 249-372, 18 figs, in text. 
This important monograph of the ticks of the world, iiiifortiinately, is obtained 

only with considerable difTiculty. 

1902.— Notes sur Ics Ixodides, I <Arcli. Parasil., vol. (i, i)p. I0!»-]2S, figs. 6. 
1904.— Notes sur Ics Ixodides, II <Ar<'h. Parasit.. vol. S ( 1904 i, no. 3. pp. 444- 

4(i4, figs. 2. 
190ri.— N",.ics sur le>< ixo<li<l(''s. III <Arch. Parasil., vol. 9 (1904), no. 2, pp. 

22.5-241 . 

NUTTALL, (<. H. F. 

1899. — Tlie role of insects, arachnids, and myriapods as carriers of human and 
animal diseases which are produced by bacteria and animal parasites 
Hyg. Rundschau, vol. 9. no. 5, pp, 209-220; no. 6, pp. 275-289; no. 8, 
pj.. 393 (OS; no. 10. ])p. 503-520; no. 12. i)p. 606-620. 

OsTiORN. 11. 

1896. Insc<-isatlVcling domestic animals V. S. I). A., Div. Eiit., bul, 5, n. s. 
Packard, A, S., Jr. 

1869.- Pisl of hymciioplcroiis an<l Icpidoplcn.us in.^ecls collccled by tlic Smith- 
sonian cxpcilitioii to South .\m<'rica, under Prof . .himes Orton; appen- 
dix to report on Articulates. .\nn. Pv('i>t. Pcabody Academy of Science, 
pp. 1-14. 

Contains descriptions of following new species of American fonns: Ixodes per punc- 
Intus, unipunctata {=Amb'y')wma americanum), leporis-palustris {Hsemaphysalis), 
chordeilis, borin (-^ Mirgircpux anvnUit-us), hihronii, vnponenxis (P.. A.), albipictns 
(X. .V,), iiifiroliiiralns. cookri. 

1873.— Descriptions of new insects. Aracluuda V . S. (W'ological Survey of 
M<.nt., Idaho, Wyo., Utah. 1872; pp. 740-741. 

Includes Ixodes bon'.i Kiley (= Mcrq-trdpiis nvmihitiis) and Argas americanus n. sp., 
with figures of (>aeh. 

Rn.EY, C. Y. 

1893. — Rei)ort on a small collection of in.-^ects made during the Death Valley 
Expedition <'TT. S. D. A., ]5iol. Surv., N. A. Fauna, no. 7, Part II, 
}). 252. 
Lists spven si)ecies of Ixodida — determined by Doctor Mar.x. 

Salmon, D. E.. and Ch. W. Stilks. 
1901. — See al)ove. 

Say, Thomas. 

1821. — An account of the arachnides of the United States <Jom-n. Acad. Nat. 
Sci., vol. 2, pp. 59-83; Leconte edition, vol. 2, pp. 9-24. 

Descriptions of the following species of Ixodes: annulalus ( Margaropus), errati- 
cus, scapularis, orbiculatus {=Amb'.yo'mma americanum?), rariabiUs (Dermacentor), 
fuscous (fuscus), crenatum (=Amblyomma cajennense), punctulatus {= Dermacentor 
variabilis f). 



BIBLIOGRAPHY. 75 

Stoll, Otto. 

1886-1893.— Arachnida Acaridea Biol. ('oiit.-Ain., pp. J8-24, i)ls. 12-14. 
Five sppcios are discussed. 

Ward, Henry B. 

1899.— The ticks of Nebraska ^Nebraska Slate Board of Aii;riculfurc, annual 
report for 1899, pj). 19.V205. Rei)rint: Studies iV.mi the Z..o]oi,ncal 
Laboratory, no. 38. 
Wheler, F,. Cx. 

1906.— British ticks Journ. .V^v. Sei., vol. 1. pp. 100 429. pis. 5-10. 
Williams, S. R. 

1905. — Anatomy of Boophihis (inniildtiis Say <,Proc. Boston Soc. Nat. Hist., 
-^ol. 32. pp. 313-334, pis. 18-22. 



XDEX 



Page. 

Amblyomnin amerirannm, bibliographic reference 67 

copulation before gaining host occasional 12 

habits and life history 59-60 

Ixodes orhiculatus a synomym (?) 74 

unipunctata a synonym 74 

outranks Margaropus annulatus as a pest, locally. ... 59 

cajennensc, habits, life history, synonyms 60-62 

complanatum=A. maculatuw 62 

habits 58 

hebraeum, bibliographic reference 70 

transmitter of heartwater in cattle, sheep, and goats. ... 58 

maculatum, habits, synonyms 62-()4 

on dog 50 

nuxtnm=A. cajennensc , 60 

multipunctum, on antelope and tapir 58 

ovatum^A. maculatum 62 

pilosum , in Galapagos Islands 73 

rubnpes=^A. macnlatum 62 

sculptus=A. cajennensc 60 

species in North America 58 

structure 54, 58 

table of species in United States 58 

tenellum^A. maculatum 62 

tigrinum^:=A. maculatum 62 

tnste^A. maculatum 62 

tuberculatum , habits and distribution 64 

in Florida 73 

Ant-eater, host of Amblyomma cajennensc 61 

Antelope. (See Dicranoccrus furcatus.) 

Antiheliotropism, in cattle tick 34 

Margaropus {Boophilus) microplus 34 

Ants, enemies of Ornithodoros moubata 37 

Arctomys monax, host of Ixodes cookei 55 

Argas americanus, bibliographic references <>!>, 73, 74 

^A. miniatus 42 

- habits 42 

miniatus. haljits, life history, synonym 42-45 

transmitter of spirillosis of fowls 42, 71 

persicus, bibliographic reference 71 

sanchczi. habits 42 

in Arizona and Now Mexico 73 

spp. , effects of bites 70 

transversa, in Galapagos Islands 73 



78 INDEX. 

Page. 

Argasida\ habits 41 

table (if genera 40 

ArinadilUdium riilgatr. eating eggs of cattle tirk 36 

Arpagostoma. ( See Ticks.) 

Ass. host of Margaropits annulatus 35, 39 

Ornithodoros megnini 45 

Bats, hosts of Ixodes ricinus 56 

Beaver, ho.st of Dermacentor alhipidus 52 

Birds, enemies of cattle tick 37, 39 

hosts of Argas 42 

Argasi(la> 41 

Ixodes ricinus 56 

sea. hosts of Coratixodes - 54 

Boophihis annulatus=Margaropus annulatus 9 

aastralis, in Philippine Islands : 69 

=;Margaropus 9 

Bufo ugiiu, host of Ambhjomma cdjennense 61 

Capybara. (See Ilydrocharrus capybara.) 

Cariacus canadensis, host of Dermacentor albipictus 51 

Cat . host of Ixodes cookei and /. ricinus 56 

Maryaropus ayinulatus 34, 68 

wild, liosl of Dermacentor bifarcutiis 50 

Ixodes ricinus 56 

Cattle, hosts of Atriblyotnuia americanum 59 

cajennense 61 

mac'ulatum 62 

Chrysomyia macellaria 12 

Dermacentor occidentalis 51 

/ variabilis 50 

Ixodes ricinus 56 

Ornithodoros megnini 45 

infestation by cattle tick affected by ration 35-36 

tick. (See Tick, cattli', and Margaropus annulatus.) 

Ceratixodes, structure and hosts 54 

Cercns cavipestris, host of Amhhjomma maculatuni 63 

Chalcidoid, with remains of cattle tick, probabh' jiarasite of dipieroii 36-37 

Chicken. {See also P'owls. ) 

liouse. occurrence of Dermacentor parumapirtus 50 

Chrysomyia macelhiriu. attack on cattle following that of tick 12 

Deer, host of Atjibh/omina americanum 59 

l>irmac<iitor nitens 52 

occidentalis 51 

l:<>d.s srupularis 57 

Margurnpus (innulatus ' -^4, 39 

Dcrmncciitor (intcriniiiuui author.'^, not Linn;vus,=7>. rnriabilis 50 

in ( 'olorado 73 

{ variabilis) Curtice^D. occidentalis 51 

(dbipiclus. distribution and hosts 51-52 

Packard (Am. Nat., and Guide;, D. rcticulatus Salmon 

and Stiles (in part) a synonym 50 

bifurralus. on wild-cat 50 

clectus^D. variabilis 49, 50 

nigroiinealus, in United States 50 



INDEX. 79 

Dermacentor nilens, distribution and hosts 52 

Occident alls, distribution, hosts, sjmonjTns 51 

Neumann, D. reliculatvs Sahnon and Stih's (in part) a 

synonym 49-50 

'^[les=^ Amblyovimo indculdtiim •)2 

transmitter of Rocky Mountain spotted fever 72 

)>(inivt(ij>ertus, in a chicken house and on man 50 

reticulutus Fabricius, not yet found in United States 50-51 

Salmon and Stiles (in part)=Z). albipictits 50 

Salmon and Stiles (in part)=i). ocddenlalis 49-50, 51 

species in United States, structure 49-50 

variabilis, habits and life history 50 

in Florida 73 

synonyms 50 

variegatus, listed from United States liy Salmon and Stiles 49 

Dicranocenis furcatus, host of Amblyomma tiwlli /unu-tuni 58 

Dipteron, probable parasite of cattle tick 37 

Dipterous larva>, enemies of cattle tick 3() 

Disease, human, transmission by ticks, bibliography 72 

possilile transmission by Aniblyonniid mjennense til 

Denuaceiilor alhipidus 52 

transmission l)y foreign ticks, bibliography G9-72 

ticks 40 

Dog, host of Ambhjonivia nmericanum 59 

cajennense 61 

maculatutn 50, 62 

Dermacentor variabilis 50 

Ixodes ricinus 56 

scapularis ' 50, 57, 58 

Mdrgaropus annulatus 34-35 

Ornithodoros viegnini 45 

Rfnpicephalus sp 47-48, 50 

Dove. (See Zenaidura macroura.) 

wild turtle, host of Argas sanchezi 42 

' • Elk . " " (See Cariacus canadensis . ) 

Feed-lot system for eradicating cattle tick ;'>7-:'.8, (57 

Felis pardalis, host of Ixodes ricinus 56 

Ferret, host of Ixodes ricinus 5() 

Fever, African coast, of cattle, transmission l)y live species of Rhi])ic('phalus. . 48, 70 

human, transmission by ticks 72 

in cattle, said to be produced l)y Ainbh/oiniiui cajennense 61 

Rocky Mountain spotted, transmission by Dermacentor occidentnlls 72 

splenetic. {See Fever, Texas. ) 

Texas, losses 1 1-12 

may it be transmitted by Ixodes ricinus? 57 

transmission by cattle tick ( Margaropus annulatus) .... 10, 11-12, 40, ()9 
" Fire-ant. ' ' (See Solenopsis geminata. ) * 
Flea, jigger. (See'Sarcopsylla penetrans.) 
Flooding. (iSee aZso Submergence. ) 

effect on seed ticks of cat th- t ick 24-25- 

Fowls, enemies of cattle tick 37, 39 

Fox, host of Ixodes cookei and /. ricinus ,,.,,..,,,, ,'!!. 56 

Freezing, (^ee Temperature.) 



80 INDEX. 

Page. 

Goat, host of Amblyomina americanum 59 

Ixodes cookei and /. ricinus 5() 

Gonixodes rostralis^Hsernaphysalis leporis-palustris 53 

Gophenis polyphemus. host of Amblyornma tuberculatum 64 

Ornithodoros furicata 4(i 

sp., host of Amblyornma tuberculatum (M 

Greasing, method of eradicating cattle ticks 07 

Guinea pig, not a host of Margaropus annulatiis 34 

Hxmaphysalis chordeilis, in United States 53 

condnna, in United States 53 

leachi, transmitter of malignant jaundice* of dogs 53 

leporis-palustris, habits, synonym 53-54 

structure and United States species 52-53 

Hand-picking, method of eradicating cattle ticks 67 

Hares, hosts of Hsemaphysalis leporis-palustris 54 

Ixodes ricinus 56 

Heartwater, in cattle, sheep, and goats, transmission l)y Amblyomma hebraeum. 58 

Hedgehog, host of Ixodes ricinus • 56 

Hog, host of Amblyomma americanum 59 

Ornithodoros turicata 46 

Horse, host of Amblyomma americanum 59 

rajennen-se 61 

maciilatum 62 

Dermaccntor nitens 52 

occidentalis 51 

Hirmaphysalis leporis-palustris 53 

Ixodes ricinus 56 

Margaropus annulatus 35, 39 

Ornithodoros megnini 45 

Host relations of ticks 12 

Houses, occurrence of Argas sanchezi 42 

Hydrochserus capybara. host of Amblyomma cajennense 61 

Insecticides, resistance of Argas miniatus • 45 

Ixodes albipictus Packard (Am. Nat. & Guide )=Dermar£ntor albipictus. 

Packard (1st Peabody Acad. Rept.)^=I)ermacentor variabilis. . . 50 

reference to original description 74 

americanus, reference to original description 73 

angustus, in United States 55 

annulatus= Margaropus annulatus 74 

reference to original description 74 

arcticus, in United States ' 55 

■ bibronii, reference to original description 74 

bifurcatus=Dermacentor bifurcatus 50 

horealis, bibliographic reference. 73 

bons, bibliographic references 74 

= Margaropus annulatus 74 

brunneus, in United States 55 

cali/orniais, in United States 55 

chordeilis. reference to original description 74 

cookei, hosts, synonyms 55-56 

reference to original description 74 

crenatum = Amblyomma cajennense 60, 74 

• reference to original description 74 



INDEX. 81 

Page. 

Ixodes cruciarius=I. cookei 55 

reference to original description 73 

dentalus, in United States 55 

diver sifossus, in New Mexico 73 

erraticus, reference to original descrii)ti(in 74 

fimbriatus^ Ixodes borealis, male 73 

frontalis, in United States 55 

fuscous ifuscus), reference to original description 74 

fuscus. in United States 55 

hal)its 55 

herrer;iR= Amblyomma americanum (iO 

hexagonus S. & S.=I. cookei 55 

var. longispinosa= I. cookei 55 

inchoatiis, in United States 55 

leporis-palustris=IIscmaphysalis leporis-palustris 74 

reference to original description 74 

naponensis, reference to original description 74 

ni(]rolineat\is=Dermacentor nigrolineatus 50, 55 

reference to original description 74 

(?) odontaLgise= , reference to original description 73 

orbiculatus= Amblyomma americanum (?) 74 

reference to original description , 74 

perpunctatus, reference to original description 74 

pilosus, bibliographic reference 71 

drying up of engorged females in captivity Ijefore ovipositing. . . 55 

plumbeus ( ?) , longevity of larvae 55 

punctidatus=Dermacentor variabilis (?) 50, 74 

reference to original description 74 

quinquestriatus=Dcrmacentor variabilis 50 

reference to original description 73 

re.duvius=I. ridnus 70 

ricinus, bibliographic references (17, 70, 72 

connection with "louping ill" of sheep only accidental 56 

habits and life history 56-57 

in Santa Cruz Mountains, California 58 

may 't transmit Texas fever in United States ? 57 

oviposition process 1(5 

transmitter of European piroplasmosis of catth^ 57, (J8 

type of genus 54 

robertsoni= Dermacentor variabilis • 50 

reference to original description 73 

scapularis, drying up of engorged females in captivity Ix'forc ovipositing. 55 

habits 57-58 

in Florida 73 

on dog 50, 57, 58 

reference to original description 74 

sculptus, distribution and host 58 

species in United States 55 

sp., from Arizona 73 

structure in genus 54 

unipunctata=Amblyo)nina aincricniinni 74 

reference to original description 74 

5795— No. 72—07 6 



82 INDEX. 

Page. 

Ixodes urise, in United States 55 

variabilis^Dermacentor variabilis 74 

reference to original description 74 

Ixodidee, subfamilies and genera 46 

table of subfamilies and genera 40 

Ixodinse, structural character and genera 46 

structure of genera 54 

table of genera 41 

Ixodoidea. (SeeTu-ks.) 

"Jackdaw." (See Quiscaliis major inacronriis.) 

Jaundice, malignant, of dogs, transmission l)y Hxmuphysnlis icarlii 53 

Jigger flea. (See Sarcopsijlla penetrans.) 
Kingbird. (See Tyrannus tyrannus.) 

Leech, use of Amblyomma maculatum ])y South American Indians. 63 

Lizards, hosts of Ixodes ricinm 56 

Llama, host of Ornithodoros turicata 46 

Lepns callotis, host of Dermacentor variabilis 50 

palnstris, host of Ilieinaphysalis leporis-paliistris 53 

sylvatieus, host of Ixodes rxeinus 56 

Longevity of Argas miniatns 44-45 

Ixodes 55 

Ornithodoros mcgnini 46 

"Loujjing ill," of slieep, connection of Ixodes ririniis only accidental o9 

Mammals, hosts of Amblyonuna americamun • 59 

Argas 42 

Argasida; 41 

Man, host of Amblyonuna americanum 59 

cajennense . 61 

maailatwn 62 

Argas 42 

Dennaeentor oecidentalis 51 

parumapertus 50 

Ixodes rieinus 56 

Margaropvs annulatus 35 

Ornithodoros megnini 45 

turieata 46 

Margaropus unnalatits. (See also Tick, cattle.) 

habitat 49 

host relations 34-35, 39 

in Florida 73 

New Mexico 73 

Ixodes bovis a synonym 74 

outranked as a pest, locally, by Amblyomma ameri- 
canum 59 

placed in genus Rhipicephalus by Neumann and Ful- 
ler 47 

transmitter of splenetic or Texas fever 40, 69 

var. argenfina, habitat 49 

aiistralis, habitat 49 

M. microphis prol)ably a synonym 49 

calcarata, habitat 49 

caudata, habitat 49 



INDEX. 83 

I'age. 

Margarnpus atiniddtiis, v;ir. (hcolortilii.s, etc., sludii's l)y Loiinslmiy 10 

luil)itat 19 

viabilit y in eggs 15- 1 (i 

correct generic nanu' for cattle tick (Boophihis anntdatus) 9 

habits and structure distinguishing it from Rliipiccphahis 47, 49 

micropliis, antiheliotropism 34 

bil)liograpliic reference 70 

genital apparatus 14 

locomotion M 

probably= M. annulatus \ar. aiialralix 49 

reattachment to host L'9 

Marmot. (See also Arctomys monax.) 

host oi Ixodes cookei 56 

Mice, hosts of Ixodes ricinus ^ 50 

probable enemies of cattle tick '.M 

Mink. (See also Putorius vison. ) 

host of Ixodes ricinus 5G 

'"Mostacilla,"' Guatemalan name for young of Anihlj/omnid cajennense (il 

Mule, host of Amhlyomma americanum 59 

Ixodes ricinus 50 

Margaropus annulatus 35, 39 

Opossum, host of Ixodes ricinus 50 

Orniihodoros americanus=0. megnini, adult 73 

habits 45 

megnini, habits 45-40 

Ornithodoros americanus and RJiijnrJiopriniii spiuosum 

synonyms 7;i 

movbata, bibliographic reference 72 

enemies 37 

transmitter of human ti<k-f('vcr 45 

'savignyi, hal)its 45 

is it identical with (). turicntu? 40 

spp. , effects of bites 70 

sp., suspected transmitter of cattle disease 45 

furicata, habits 40 

is it identical with (). sarignyi.' 4(i 

Otter, host of Ixodes cookei 50 

Oviposition process in ticks 10-17 

Panther, host of Ixodes ricinus 50 

Paradoxurus, host of Hsemaphysalis leporis-palustris 54 

Pasture-rotation system, for eradicating cattle tick :W, 07 

Peccary, host of Amhlyomma cajennense 01 

Phoridse, a species bred from cattle tick '. . . 30 

Picketing, method of eradicating cattle ticks. {See Tie-roi)e. i 

Pigeons, hosts of Argas 42 

Piroplasma higeminum, found in both Europe and America .' 57 

Piroplasmosis, European, of cattle, transmission by Ixodes ricinus 57, 08 

Polecats, hosts of Ixodes ricinus 56 

Porcupine, hosts of Ixodes cookei 50 

record as host of cattle tick pr()l)ably erroneous 35 

Practical application of information 37-39 

Prairie dogs, hosts of Ixodes scapularis 58 



84 INDEX. 



"Prootoquie," term used by Lahille for period, in tick, previous to oviposition. . 14 

Pittoriiis lison, host oi Ixodes cookei 56 

Quail, host of Argas sanchezi 42 

Quiscalus viajor macrourus, enemy of cattle tick 'M 

Rabbit. (See also Lepus callotis, L. palustris, and L. sylvaticus.\ 

host of Hxmaphysalin leporis-palustris 53-54 

Ixodes ricinus 56 

Margaropus annulatus M 

rc^cord as host of cattle tick probably erroneous 35 

Raccoon, host of Ixodes cookei 56 

Rats, enemies of Ornithodoros moiibata 37 

Rlii])ic('pluilin:e, structural character and genera 46 

tal>le of genera 40-41 

Rhipkephaliis uppendiadatus, bibliographic reference 71 

bursa americamis, reported from Jamaica 47 

capensis, bibliographic reference 71 

decoloratus, transmitter of spirillosis in cattle 72 

evertsi, bibliographic reference 71 

habits and structure distinguishing it from Margaropus 47, 49 

sanguineus (Ann. Rpt. Bur. Animal Industry f. 1905, p. 35 1 = 

Rhipicephalns sp. of j)resent bulletin 48 

in Panama 47 

simus, bibliographic reference 71 

sp., habits and life history ^. . . '. 47-49 

in Colorado 73 

on dog 50 

same as R. sanguineus in Ann. Il])t. liur. Animal Industry 1. 

1905, p. 35. 48 

undetermined species from Colorado and Porto Rico 47 

Rhynchoprium spinosum=Ornithodoros megnini. nymph 46, 73 

Roebuck, host of Ixodes ricinus 56 

Sarcopsylla penetrans, introduction into Soutli Africa 46 

Screw-worm fly. (See Chrysomyia macellaria.i 

Sheep, host of Dermacentor occidentalis 51 

Ixodes cookei and /. ricinus 56 

Margaropus annulatus 35, 59 

Ornithodoros megnini 45 

Skunk, host of Ixodes cookei 56 

Snake, Ornithodoros turicata in burrow 46 

Soiling system, for eradicating cattle tick. {See Feed-lot system. ) 

Solenopsis geminata, enemy of cattle tick 36 

Sp(>rmophile, host of It odes cookei 56 

S})irillosis of fowls, transmission by Argas miniatus 42 

Spirillum (Spirachsete) obermeien, prol)able cause of human tick fever 45 

Splenetic fever. {See Fever, Texas.) 

Squirrel, host of Amblyomma americanum 59 

Ixodes ricinus 56 

Stag, host of Ixodes ridnus 56 

Steer, used in experiments on cattle tick 27 

Submergence, in water, effect on engorged adult cattle ticks 32-33, 39 

incubation of cattle tick 22, 39 

Sunlight, direct, effect on adult cattle ticks ; 32 

Tapir, host of Amblyomma nrultipunctum 58 



INDEX. 85 

rage. 

Temperal ufc. effort on engorged females of cattle t ick 31-32 

seed ticks of cattle t ick 24-25 

in relation to incubation of cattle lick 19-21 

Texas fever. [See Fever, Texas. ) 
Tick, adobe. (See Argas sanchezi.) 

Australian. {See Boophilus australis.) 
black-legged. (See Ixodes scapularis.) 

blue, common, bibliographic reference 69 

bont. (See Amblyomvia hebrseum.) 
castor bean, American. (See Ixodes cookei.) 
European. (See Ixodes ricimis.) 
cattle. (See also Margaropus annidatus.) 

adult female, description 31 

male, description 30 

stage 30-34 

adults as affected l)y direct sunlight 32 

submergence in water 32-33, 39 

attachments of specimens to each other 35 

bibliography ()5-()9 

control. (See eradication methods.) 

descriptions of adults 30-31 

development as affected by ration of host 35-36 

on host 28-30 

dropping from host 33 

egg stage 15-23 

eggs as affected by heat and cold 21-22 

submergence in water 22, 39 

indication of viability 15-16 

number deposited 17 

percentage hatching 23 

engorged adults as affected by submergence in water 32-33, 39 

continuous cold and heat 31-32 

enemies 3()-37, 39 

eradication, from United States possible 9 

nicthods 37-39 

most important factor 13 

host relations - 34-35, 39 

importance of knowledge of life history 9-10 

incubation 17-23 

period 17-21 

infestation affected by ration of host 35-36 

knowledge of variations of periods in life history esscmtial 13 

lack in knowledge of local climatic variations and dissemination . 10 

larval stage 23-30 

nonparasitic period 23-2() 

parasitic period 27-30 

life history 13-33 

locomotion 33-34, 39 

longevity of secnl ticks 25-26 

losses 11-12 

male, position on host with relation tf) female 30 

molts of larval stages 28-29 

most important factor from standpoint of control 13 



86 INDEX. 

Pasre. 

Tick, cattle, nymphal stage 28-29 

oN'iposition period 14-15 

process 16-17 

periods in life history upon which means of control are based. . 38 

practical application of data 37-39 

preoviposition period 14 

previous work on life history 10 

reattachment to host 29-30 

seed ticks as affected by submergence in water 24-25, 39 

longevity 25-26 

tick stage 23-30 

nonparasitic period 23-26 

parasitic period 27-30 

submergence as affecting eggs 22, 39 

engorged adults 32-33, 39 

sunlight as affecting adults 32 

temperature as affecting engorged females 31-32 

incubation 19-21 

seed ticks 24-25 

transmitter of Texas fever 10, 11-12 

Cayenne. (See Amblyomvia cajennense.) 
dog, American. (See Dermacentor variabilis.) 

brown. (See Rhipicephalus sp.) 
elk. (See Dermacentor albipictus.) 

fever, human, bibliographic reference 72 

t ransmission by Ornithodoros moubata 45 

Norlh American. (See Margaropiin annnlatus and Tick, cattle. ) 
fowl. (S(>c also .l/Y/r;.s' miniatus.) 

bibliographic reference 70 

Gulf coast. (Se(> Amblyomma maculalum.) 
horse, tropical. {See Dermacentor nitens.) 
land turtle. (See Amblyomma tuberculatum..) 
lon(> star. (See Amblyomma americanum.) 
net. (See Dermacentor occidentalis.) 
paralysis. (See Ixodes pilosus.) 
rabbit. (See Hsemaphysalis leporis-palustris.) 
s(;ulptiu'ed. (See Ixodes sculptus.) 
spinose ear. (See Ornithodoros megnini.) 
turicata. (See Ornithodoros turicata.) 
wood. (Se(> Dermacentor variabilis.) 

Ticks, l)ibliography 65-75 

castor bean . ( See I xod es . ) 

classification and distribution, bibliography 73-75 

habits 40-64 

foreign, that transmit disease, bibliography 69-72 

graphic table for separation of families and genera, from Lahille 41 

habits 40-64 

host relations 12 

key to families, subfamilies, and North American genera 40-41 

• life history, general statement 12-13 

transmitters of disease 40. 65-72 

work by Lahille 10 

"yearling," name given to ticks in nymphal stage 13 



INDEX. 87 

Page. 

Tie-rope, method of eradicating cattle ticks 67 

Toad. {See Bufo agua.) 

Turtle, land. (See Gopherus ijolypheriias.) 

Two-field method of eradicating cattle ticks 67 

Wapiti. (See Cariacus canadensis.) 

Weasel, host of Ixodes cookei 56 

Wolf, host of Amblyomma americaniun 59 

Woodchuck. (See Marmot and Arctomys monar.) 
Xerobates polyphemus. (See Gopherus polypheinus. ) 

"Yearling ticks," name given to ticks in nymphal stage 13 

Zenaidura macroiira, host of Argas sanchezi 42 



F. Mr '08 



^ 



